Ultrastructural changes of the midgut epithelium in Isohypsibius granulifer granulifer Thulin, 1928 (Tardigrada: Eutardigrada) during oogenesis

The midgut epithelium of Isohypsibius granulifer granulifer (Eutardigrada) is composed of columnar digestive cells. At its anterior end, a group of cells with cytoplasm which differs from the cytoplasm of digestive cells is present. Probably, those cells respond to crescent-like cells (midgut regenerative cells) described for some tardigrade species. Their mitotic divisions have not been observed. We analyzed the ultrastructure of midgut digestive cells in relation to five different stages of oogenesis (previtellogenesis, beginning of the vitellogenesis, vitellogenesis—early choriogenesis, vitellogenesis—middle choriogenesis, late choriogenesis). In the midgut epithelium cells, the gradual accumulation of glycogen granules, lipid droplets and structures of varying electron density occurs. During vitellogenesis and choriogenesis, in the cytoplasm of midgut cells we observed the increasing number of organelles which are responsible for the intensive synthesis of lipids, proteins and saccharides such as cisterns of endoplasmic reticulum and Golgi complexes. At the end of oogenesis, autophagy also intensifies in midgut epithelial cells, which is probably caused by the great amount of reserve material. Midgut epithelium of analyzed species takes part in the yolk precursor synthesis.     

Comparative midgut ultrastructure of unfed larvae and adult mites of Platytrombidium fasciatum (C.L. Koch, 1836) and Camerotrombidium pexatum (C.L. Koch, 1837) (Acariformes: Microtrombidiidae)

The midgut of unfed larvae and adult mites of Platytrombidium fasciatum (C.L. Koch, 1836) and Camerotrombidium pexatum (C.L. Koch, 1937) (Acariformes: Microtrombidiidae) was investigated by electron microscopy. The sac-like midgut occupies the entire body volume, ends blindly and is not divided into functionally differentiated diverticula or caeca. The midgut walls are composed of one type of digestive cell that greatly varies in shape and size. In larvae, the lumen of the midgut is poorly recognizable and its epithelium is loosely organized, although yolk granules are already utilized. In adults, the midgut forms compartments as a result of deep folds of the midgut walls, and the lumen is well distinguished. The epithelium is composed of flat, prismatic or club-like cells, which may contain nutritional vacuoles and residual bodies in various proportions that depend on digestive stages. In both larvae and adult mites, parts of cells may detach from the epithelium and float within the lumen. The cells contain a system of tubules and vesicles of a trans-Golgi network, whereas the apical surface forms microvilli as well as pinocytotic pits and vesicles. Lysosome-like bodies, lipid inclusions and some amount of glycogen particles are also present in the digestive cells. Spherites (concretions) are not found to be a constant component of the digestive cells and in adult mites occur for the most parts in the midgut lumen.     

Ultrastructure of the digestive system and the fate of midgut during embryonic development in Porcellio scaber (Crustacea: Isopoda)

Microscopic anatomy of the digestive system in embryos and larvae of the terrestrial isopod crustacean Porcellio scaber was investigated by light bright field, fluorescence and electron microscopy. During marsupial ontogenetic development the event-dependent staging was used to discriminate the various embryonic stages. At the late embryo stage the differentiation of the ectodermal part of the gut into the complex filtering foregut and the hindgut with absorptive and transporting functions is accomplished. The gut of the marsupial manca larva is fully developed and similar to that of the adult. In early embryos the endodermal midgut gland primordia are filled with yolk and lipid globules. In late embryos the epithelium of paired midgut gland tubes is composed of two cell types; one of them exhibits orange autofluorescence. The endodermal cells located between the foregut and the midgut glands of late embryos form the prospective midgut. The cells have electron dense cytoplasm, abundant glycogen fields, endoplasmic reticulum, dictyosomes and numerous vesicles. In the adults the endodermal cells of the midgut remain only in the midgut gland ducts which connect the midgut glands and the foregut. Details of the cellular ultrastructure and morphogenesis of the ectodermal and endodermal parts of the digestive system during embryonic development of Porcellio scaber provide data for further phylogenetic and comparative studies in peracaridan crustaceans and other arthropods.     

The digestive system of the fish ectoparasite Dolops ranarum (Crustacea: Branchiura)

Reconstructions from serial sections reveal that the digestive system consists of a pharynx esophagus, crop, midgut, and rectum. Two main stems from the branched enteral diverticula are connected to the lateral regions of the crop by right rind left arms. Glandular tissue surrounds the enteral diverticula proximally. A strong sphincter separates the crop and midgut, whereas the midgut and rectum are separated by a weak sphincter. Cuboidal epithelium lines the pharynx, esophagus, crop, and rectum, whereas cuboidal and club-shaped epithelia line the midgut. The cuboidal cells possess elongated nuclei and numerous vacuoles, suggesting that absorption takes place in the midgut.     

Autophagy as the cell survival in response to a microsporidian infection of the midgut epithelium of Isohypsibius granulifer granulifer (Eutardigrada: Hypsibiidae)

The midgut epithelial cells of many invertebrates may possess microorganisms which act as symbionts or pathogens (bacteria, microsporidia, viruses). During our previous studies on Isohypsibius granulifer granulifer Thulin, 1928 (Tardigrada, Eutardigrada), which examined alterations of the midgut epithelium during oogenesis, we found that some of the specimens were infected with microsporidia. All stages of pathogens occurred in the cytoplasm of the digestive cells in the midgut epithelium of I. g. granulifer that were infected with microsporidia: meronts, sporonts, sporoblasts, and spores. The cytoplasm of the digestive cells was rich in mitochondria, cisterns of rough endoplasmic reticulum (RER), and Golgi complexes. Autophagy in the digestive cells of the dorsal midgut was much more intensive in comparison with noninfected specimens. Membranes of phagophores surrounded the pathogens forming autophagosomes. These latter structures fused with lysosomes forming autolysosomes and residual bodies appeared. Neither glycogen granules nor droplets of varying electron density, which accumulated in digestive cells during vitellogenesis and choriogenesis, appeared in individuals with microsporidia. While the midgut epithelium in noninfected specimens takes part in vitellogenesis and choriogenesis, in infected specimens, midgut cells are involved in the process of autophagy as a survival strategy.     

The fine structure of the midgut epithelium in a centipede,Scolopendra cingulata (Chilopoda,Scolopendridae), with the special emphasis on epithelial regeneration

Scolopendra cingulata has a tube-shaped digestive system that is divided into three distinct regions: fore-, mid- and hindgut. The midgut is lined with a pseudostratified columnar epithelium which is composed of digestive, secretory and regenerative cells. Hemocytes also appear between the digestive cells of the midgut epithelium. The ultrastructure of three types of epithelial cells and hemocytes of the midgut has been described with the special emphasis on the role of regenerative cells in the protection of midgut epithelium. The process of midgut epithelium regeneration proceeds due to the ability of regenerative cells to proliferate and differentiate according to a circadian rhythm. The regenerative cells serve as unipotent stem cells that divide in an asymmetric manner.Additionally, two types of hemocytes have been distinguished among midgut epithelial cells. They enter the midgut epithelium from the body cavity. Because of the fact that numerous microorganisms occur in the cytoplasm of midgut epithelial cells, we discuss the role of hemocytes in elimination of pathogens from the midgut epithelium. The studies were conducted with the use of transmission electron microscope and immunofluorescent methods.     

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.     

Ultrastructural changes in the midgut epithelium of the first larva of Allacma fusca (Insecta, Collembola, Symphypleona)

Abstract. In the newly hatched larva in Allacma fusca , the midgut epithelium was fully developed and formed by flattened epithelial cells surrounding the yolk mass in the midgut lumen. Immediately after hatching, the first larva began to feed; the migut lumen was filled with the yolk mass and food (mainly algae). Regenerative cells typical of the developing midgut epithelium of many insects were not observed. Initially, midgut cells of the larva were cuboidal but became columnar in shape with distinct regionalization in the distribution of cell organelles. Furthermore, urospherites appeared in the midgut cell cytoplasm, i.e., structures characteristic for the midgut epithelium of insects having no Malpighian tubules. As a result, cells with the capacity for digestion, absorption, and excretion were observed to be completely formed in the first larval stage.     

Morphofunctional changes in the midgut of tick females of the genus Ixodes (Acari: Ixodidae) during and after feeding

Cyclic changes of the midgut epithelium were observed in females of 5 ticks species of the genus Ixodes during 7-10 days of feeding. The midgut epithelium of unfed females is represented by the digestive cells of nymphal phase and stem cells. The digestive cells of nymphal phase are functional during 1.5-2 days after attachment of the tick, and then, after the tearing away they go into the gut lumen. The secretory cells substitute the digestive cells of nymphal phase and finish their growth during the 4-4.5 days. Secretion of digestive enzymes is performed by the holocrine type with tearing away a whole cell. Intracellular digestion takes place in the digestive cells of four consequent generations. The secretory and digestive cells form a peritrophic matrix on their surface. The presence of peritrophic matrix gives an evidence the maturity and functional activity of the secretory and digestive cells. We suggest, that the peritrophic matrix takes part in intracellular digestion, namely in the process of micropinocytosis. The phagocytosis was not found in the ticks investigated. Digestion in the midgut lumen is performed by enzymes of the ruptured secretory and digestive cells, that is proved by the haemolysis of erythrocytes in the zone of their contact with these cells. The digestive cells of each generation functioned almost synchronously, with largest difference in starting about 12 hours.     

Histological studies on the development of the digestive system of the clownfish Amphiprion percula and the time of weaning

This study investigates the histological development of the digestive system of the common clownfish, Amphiprion percula Lacépède 1802, as a means to predict the time when the early juveniles can begin to digest inert feeds. The incubation period is completed within 7 days. At the time of hatching the alimentary canal is advanced and larvae start exogenous feeding immediately. Three days after hatch (DAH) the yolk sac is completely absorbed and at 5 DAH small supranuclear inclusions appear in the hindgut epithelium, suggesting pinocytotic digestion. At 7 DAH gastric glands are established in the epithelium of the stomach and by 9 DAH supranuclear inclusion vacuoles have appeared in the epithelium of the midgut, indicating extracellular digestion and absorption across the lumen. It was concluded that A. percula could only effectively digest artificial feeds from 9 DAH.     

Ultrastructure of the digestive tract of Diacyclops thomasi (Cyclopoida, Copepoda) during different stages of encystment during a summer diapause

Baud, A., Cuoc, C. and Alekseev, V. 222_. Ultrastructure of the digestive tract of Diacyclops thomasi (Cyclopoida, Copepoda) during different stages of encystment during a summer diapause. — Acta Zoologica (Stockholm) 85 : 181–189 Diacyclops thomasi ( Forbes, 1882 ) has a life cycle comprising a summer diapause with whole‐body encystment at the copepodid IV stage. Formed progressively, the cyst can indicate the extent of entry into diapause. In this study the ultrastructure of the midgut epithelium during encystment of CIV was compared with that of active CIV and females. In active individuals two well‐differentiated epithelial cell types were observed: vacuolar cells (B‐cells), and cells without vacuoles but with more densely packed microvilli and abundant rough endoplasmic reticulum and mitochondria (F/R cells). In encysted specimens a striking transformation of the midgut epithelial layer was noted. Only inactive nuclei with highly concentrated euchromatin and encircled by a thin rim of hyaloplasm were observed. Compartments of both urosome and especially cephalothorax displayed large lipid‐rich lacunae. At the beginning of encystment, the midgut epithelium showed an intermediate state. Changes in ultrastructure observed in the midgut epithelium of D. thomasi clearly reflect different stages of diapause, and represent clear evidence of profound reorganization, which is progressively induced by diapause in the organism.     

Fine structure of the midgut epithelium in the millipede <Emphasis Type="Italic">Telodeinopus aoutii</Emphasis> (Myriapoda,Diplopoda) with special emphasis on epithelial regeneration

The midgut of millipedes is composed of a simple epithelium that rests on a basal lamina, which is surrounded by visceral muscles and hepatic cells. As the material for our studies, we chose Telodeinopus aoutii (Demange, 1971) (Kenyan millipede) (Diplopoda, Spirostreptida), which lives in the rain forests of Central Africa. This commonly reared species is easy to obtain from local breeders and easy to culture in the laboratory. During our studies, we used transmission and scanning electron microscopes and light and fluorescent microscopes. The midgut epithelium of the species examined here shares similarities to the structure of the millipedes analyzed to date. The midgut epithelium is composed of three types of cells—digestive, secretory, and regenerative cells. Evidence of three types of secretion have been observed in the midgut epithelium: merocrine, apocrine, and microapocrine secretion. The regenerative cells of the midgut epithelium in millipedes fulfill the role of midgut stem cells because of their main functions: self-renewal (the ability to divide mitotically and to maintain in an undifferentiated state) and potency (ability to differentiate into digestive cells). We also confirmed that spot desmosomes are common intercellular junctions between the regenerative and digestive cells in millipedes.     

Ultrastructural and cytochemical aspects of metamorphosis in the midgut of Apis mellifera L. (Hymenoptera: Apidae: Apinae)

The midgut of Apis mellifera is remodeled during metamorphosis. The epithelium and, to a lesser extent, the muscular sheath degenerate between the end of the last larval instar and the onset of pupation (prepupa). The larval epithelium is shed to the midgut lumen and digested, while a new epithelium is reconstructed from larval regenerative cells. During pupation, some reorganization still occurs, mainly in brown-eyed pupae. In pharate adult, the midgut wall shows the characteristics of adult, although some cells have pycnotic nuclei. The localization of alkaline and acid phosphatases showed that these enzymes were not involved in the reabsorption of the midgut wall.     

Electron microscopic study of the anterior midgut in Cenocorixa bifida Hung. (Hemiptera: Corixidae) with reference to its secretory function

The epithelium of anterior midgut of adult Cenocorixa bifida was examined with light and electron microscopy. The folded epithelium is composed of tall columnar cells extending to the lumen, differentiating dark and light cells with interdigitating apices and regenerative basal cells in the nidi surrounded by villiform ridges that penetrate deeply into the epithelium. The columnar cells display microvilli at their luminal surface. Microvilli lined intercellular spaces and basal plasma membrane infoldings are associated with mitochondria. These ultrastructural features suggest their role in absorption of electrolytes and nutrients from the midgut lumen. The columnar cells contain large oval nuclei with prominent nucleoli. Their cytoplasm is rich in rough endoplasmic reticulum, Golgi complexes and electron-dense secretory granules indicating that they are also engaged in synthesis of digestive enzymes. The presence of secretory granules in close proximity of the apical plasma membrane suggests the release of secretion is by exocytosis. The presence of degenerating cells containing secretory granules at the luminal surface and the occurance of empty vesicles and cell fragments in the lumen are consistent with the holocrine secretion of digestive enzymes. Apical extrusions of columnar cells filled with fine granular material are most likely formed in response to the lack of food in the midgut. The presence of laminated concretions in the cytoplasm is indicative of storageexcretion of surplus minerals. The peritrophic membrane is absent from the midgut of C. bifida.     

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.     

意大利蜜蜂胚后发育过程中中肠上皮组织细胞的更替

中肠是昆虫消化、 吸收营养物质的主要部位。本研究通过形态解剖、 BrdU免疫组织化学和原位末端转移酶标记(TUNEL)细胞凋亡检测等技术, 对意大利蜜蜂Apis mellifera ligustica中肠胚后发育过程中细胞的增殖和凋亡模式进行了比较研究。结果表明：意大利蜜蜂幼虫发育早期, 中肠的增加主要来自于上皮细胞的分裂以及再生细胞的增殖。在变态发育期间, 中肠上皮经历了广泛的重组, 由再生细胞重新形成的蛹上皮替代了幼虫上皮。再生细胞在蜜蜂中肠的整个发育阶段始终存在, 为中肠的生长和更替提供了主要的细胞来源。本研究为昆虫组织细胞自噬和凋亡机制的研究提供一定的证据。     

仿刺参消化系统的组织学和组织化学研究

用组织学和组织化学方法研究了仿刺参的消化系统。消化道管壁由粘膜层、粘膜下层、肌层和外膜组成。粘膜层为假复层或单层的柱状细胞或立方细胞与粘液细胞。粘液细胞分布于前肠的前段和排泄腔。前肠和中肠上皮具蛋白酶、脂酶和非特异性酯酶活性。中肠上皮细胞游离端有密集微绒毛,游离端质膜呈碱性磷酸酶活性,上皮下有丰富的血窦,表明具吸收作用。     

Morphology and ultrastructure of the midgut in Piscicola geometra (Annelida, Hirudinea)

This paper presents information on the organization of the midgut and its epithelium ultrastructure in juvenile and adult specimens of Piscicola geometra (Annelida, Hirudinea), a species which is a widespread ectoparasite found on the body and gills and in the mouth of many types of fish. The analysis of juvenile nonfeeding specimens helped in the explanation of all alterations in the midgut epithelium which are connected with digestion. The endodermal portion (midgut) of the digestive system is composed of four regions: the esophagus, the crop, the posterior crop caecum, and the intestine. Their epithelia are formed by flat, cuboidal, or columnar digestive cells; however, single small cells which do not contact the midgut lumen were also observed. The ultrastructure of all of the regions of the midgut are described and discussed with a special emphasis on their functions in the digestion of blood. In P. geometra, the part of the midgut that is devoid of microvilli is responsible for the accumulation of blood, while the epithelium of the remaining part of the midgut, which has a distinct regionalization in the distribution of organelles, plays a role in its absorption and secretion. Glycogen granules in the intestinal epithelium indicate its role in the accumulation of sugar. The comparison of the ultrastructure of midgut epithelium in juvenile and adult specimens suggests that electron-dense granules observed in the apical cytoplasm of digestive cells take part in enzyme accumulation. Numerous microorganisms were observed in the mycetome, which is composed of two large oval diverticles that connect with the esophagus via thin ducts. Similar microorganisms also occurred in the cytoplasm of the epithelium in the esophagus, the crop, the intestine, and in their lumen. Microorganisms were observed both in fed adult and unfed juvenile specimens of P. geometra, which strongly suggests that vertical transmission occurs from parent to offspring.     

Fine structure of the midgut epithelium of Nicoletia phytophila Gervais, 1844 (Zygentoma: Nicoletiidae: Nicoletiinae) with special emphasis on its degeneration

The midgut epithelium of Nicoletia phytophila is composed of columnar digestive cells and regenerative cells that form regenerative nests. The cytoplasm of midgut epithelial cells shows typical regionalization in organelle distribution. Two types of regenerative cells have been distinguished: cells which are able to divide intensively and cells which differentiate. Spot desmosomes have been observed between neighboring regenerative cells. The occurrence of intercellular junctions is discussed. The midgut epithelium degenerates both in an apoptotic and necrotic way. Necrosis proceeds during each molting period (cyclic manner), while apoptosis occurs between each molting, when the midgut epithelium is responsible for e.g. digestion. These processes of epithelium degeneration are described at the ultrastructural level. Our studies not only add new information about fine structure of the midgut epithelium of N. phytophila, but contribute to resolving the relationships within the Zygentoma. There are no doubts about the very close sister position of Nicoletiidae and Ateluridae. The midgut epithelium characters confirm their close relationship. However we do not recommend classifying the atelurid genera only within Nicoletiidae: Nicoletiinae.     

The role of autophagy in the midgut epithelium of Eubranchipus grubii (Crustacea,Branchiopoda, Anostraca)

Eubranchipus grubii (Crustacea, Branchiopoda, Anostraca) is an omnivorous filter feeder whose life span lasts no more than 12 weeks. Adult males and females of E. grubii were used for ultrastructural studies of the midgut epithelium and an analysis of autophagy. The midgut epithelium is formed by columnar digestive cells and no regenerative cells were observed. A distinct regionalization in the distribution of organelles appears – basal, perinuclear and apical regions were distinguished. No differences in the ultrastructure of digestive cells were observed between males and females. Autophagic disintegration of organelles occurs throughout the midgut epithelium. Degenerated organelles accumulate in the neighborhood of Golgi complexes, and these complexes presumably take part in phagophore and autophagosome formation. In some cases, the phagophore also surrounds small autophagosomes, which had appeared earlier. Fusion of autophagosomes and lysosomes was not observed, but lysosomes are enclosed during autophagosome formation. Autophagosomes and autolysosomes are discharged into the midgut lumen due to apocrine secretion. Autophagy plays a role in cell survival by protecting the cell from cell death.