Observations on the integument of the water mite Arrenurus major (Acari: Parasitengona)

A study of the integument of the aquatic mite Arrenurus major Marshall is presented. When the cuticle is examined with the unaided eye and the light microscope, it appears to possess numerous tiny pits. However, scanning electron micrographs of the cuticle reveal that it is a solid surface with topographical sculpturing of the epicuticle, indicating that the “pits” are an internal phenomenon. In cuticle which has been sectioned, areas devoid of cuticular material beneath the thin exocuticle are revealed. These areas are the pits which are goblet-shaped. The integument consists of five major strata. These are from the outside to the inside: (1) a superficial layer with a maximum observed thickness of 725 Å, (2) an epicuticle with a thickness of about 900 Å and composed of at least four sublayers, (3) an exocuticle with a thickness of about 1.5 Å. Fibers of the exocuticle are arranged in a Bouligand pattern and exhibit a regularly occurring discontinuity with a spacing of 200 Å. (4) An endocuticle ranging from 15 to 20 μ in thickness. The endocuticle is characterized by bandings which superficially resemble the lamellae of insects but are not homologous, microfibers which exhibit a preferred orientation, and the presence of the pits; and (5) an epidermis lying beneath the endocuticle and extending into the pits. Pore canals are present only in the exocuticle and have their origin at the apices of the pits. The pore canals contain a central filament, and a plug is present just beneath the epicuticle.     

The structure of the ostracode carapace

The carapace ultrastructure of three Recent [Heterocypris incongruens (Ramdohr), Cypridopsis uidua (Müller), and Conchoecia belgica Müller] and one Cretaceous ostracode species [Cypridea propunctata Sylvester-Bradley] has been studied by means of standard electron microscope techniques. The carapace is shown to be an organic structure divisible into an outer epicuticle, a median exocuticle and an inner endocuticle in the three cypridacean ostracodes examined. In Conchoecia belgica only an outer epicuticle and an inner endocuticle were observed, but this may be due to preservation. The chitin structure of the carapace of calcareous ostracodes is shown to be essentially an interlocking lattice, more coarsely developed in the exocuticle; layered (lamellar) chitin is only present in the selvage and in the connective tissue joining the valves at the hinge. The non-calcareous Conchoecia belgica possesses the layered chitin structure common to other crustaceans and to insects. The relationship of the carapace structure to that present in the decapod and insect exoskeleton is discussed. The technical problems met with during this study are considered in order to outline the difficulties associated with the study of organisms of this size.     

Fine structure of the cuticle of the desert scorpion, Hadrurus arizonensis.

The structure of the sclerite and intersegmental cuticle of the opithosoma of the desert scorpion, Hadrurus arizonensis, has been examined by transmission electron microscopy. The sclerite cuticle contains a four-layered epicuticle, a hyaline exocuticle, an inner exocuticle and an endocuticle. The outer part of the hyaline exocuticle and the whole of the inner exocuticle are constructed of helicoidally arranged planes of microfibrils. Within the endocuticle, the overall architecture is not helicoidal as previously assumed, but consists of bundles of microfibrils oriented horizontally and vertically. Microbibrils of the inner exocuticle and the endocutile are seen as simple unstained rods, but those of the hyaline exocuticle are electron dense rods with an unstained central core. The intersegmental cuticle contains a four-layered epicuticle and a procuticle. In detail, its fine structure differs in most respects from that of the sclerite cuticle. Electron microscopy reveals that hyaline exocuticle, previously assumed to be continuous from sclerite to intersegmental membrane, is absent in the latter.     

中华绒螯蟹不同生理阶段腹肢的结构变化及粘液腺的发育特征

用显微及亚显微方法研究了中华绒螯蟹雌体不同生理阶段(幼蟹、未成熟蟹、成熟蟹、抱卵蟹和流产蟹)腹肢的体壁结构变化和粘液腺发育特征。体壁的上皮细胞层常与粘液腺相连,粘液腺分泌物经导管穿过各层角膜排出。不同生理状况下中华绒螯蟹的腹肢体壁各角膜层结构的比例及致密度有明显的差异,粘液腺细胞及导管的数量不同,粘液腺与上皮细胞的连接程度也不同。粘液腺是否正常分泌、刚毛囊的开闭均与中华绒螯蟹的胚胎附着有关。比较研究发现,太湖抱卵蟹腹肢体腔内腺体比温州本地抱卵蟹腹肢内腺体发达,分泌的粘液也特别多。分析认为中华绒螯蟹腹肢组织结构、粘液腺的发育和分泌状况与胚胎流产有密切的关系。     

Structure,molting, and mineralization of the dorsal ossicle complex in the gastric mill of the blue crab,Callinectes sapidus

This study examined the mesocardiac and urocardiac ossicles in the gastric mill of the blue crab to describe its structure, mineralization, and dynamics throughout the molt cycle, and to assess its possible utility in age determination. Morphologically, the mineralized ossicles are similar to the calcified dorsal carapace having a lamellate structure comprised of sheets of chitin/protein fibrils. Staining with acridine orange showed the same arrangement of an epicuticle, exocuticle, and endocuticle. In much of the mesocardiac and urocardiac ossicles, the endocuticle is very reduced, with the exocuticle predominating; the reverse of the dimensions of the exoskeleton. The lamellate structure of the ossicles was confirmed with scanning electron microscopy; however, elemental mapping by energy‐dispersive analysis of X‐rays revealed that the ossicles are mineralized with calcium phosphate, in contrast to the calcium carbonate biomineral of the exoskeleton. The medial tooth of the urocardiac ossicle is not calcified, but the epicuticle is highly elaborated and impregnated with silica. Histological examination of the ossicles demonstrated that they are molted during ecdysis, so despite the appearance of bands in the mesocardiac ossicle, it is difficult to hypothesize how the bands could represent a record of chronological age. J. Morphol. 276:1358–1367, 2015. © 2015 Wiley Periodicals, Inc.     

The Structure and Mineralization of the Carapace of the Crab (Cancer pagurus L.) 2. The exocuticle

The exocuticle of the dorsal carapace of the intermoult Cancer pagurus was studied by means of light microscopy, electron microscopy, and contact microradiography. In the exocuticle a 5-15 μm wide highly mineralized external zone was seen where prisms were not present. The cross-sectional shape and diameter of the prisms varied. The prisms were separated by interprismatic areas varying in width and in degree of mineralization. With increasing width of the interprismatic areas the diameter of the prisms decreased. The prism-less external zone may be the result of complete mineralization of the prisms. The pore canals of the exocuticle were circular in cross section and present both within the prisms and within the interprismatic areas. The lamellar system of the exocuticle was built up by layers of horizontal fibrils which were interconnected by vertical or oblique fibrils. The minerals of the external prism-less zone occurred as aggregates of crystalline matter or as large brick-like structures. Crystal edge lengths up to 300 nm were measured. In well mineralized interprismatic areas aggregates of crystals and relatively large crystals were observed. In poorly mineralized interprismatic areas small plate-like crystals occurred. Occasionally a prism-less zone was seen near the exocuticle/endocuticle junction.     

Ultrastructure and cuticle formation of the carapace in the myodocopan ostracod exemplified by Euphilomedes japonica (Crustacea: Ostracoda)

The ultrastructure and formation of the cuticle of a myodocopan ostracod, Euphilomedes japonica, are investigated utilizing scanning and transmission electron microscopy. The outer lamella cuticle consists of four layers; epicuticle, exocuticle, endocuticle, and membranous layer like in the cuticle of other arthropods. The exocuticle and endocuticle are well-calcified and the organic matrix develops within the both cuticles. The outermost layer of new cuticle (epicuticle) is secreted first and the inner layers (exocuticle, endocuticle and membranous layer) are added proximally in the pre-, and postmoult stages. The calcification takes place in the whole area of carapace at the same time together with the synthesis of organic matrix within the endocuticle. This study demonstrates that the ultrastructure and formation of the cuticle in myodocopans are different from those in podocopans, and that the myodocopan carapaces have achieved a structural diversity for adaptation to different lifestyles.     

Changes in the structure of the cuticle of Ixodes ricinus L. 1758 (Acari, Ixodidae) during feeding

Female Ixodes ricinus (ticks) are able to consume several times as much as their own weight in food. The body volume increases without the animal moulting. Special structural and material characteristics of the cuticular exoskeleton render this phenomenon possible. Only the alloscutum, which has a very pronounced lamellar structure, extends. The individual lamellae correspond to surface folds in the outer endocuticle and the epicuticle covering it. The degree of folding is correlated to the nutritional state of the animal, which can be differentiated into three significant phases. The phase-dependent morphology of the surface structure and the cuticular layers of the alloscutum are described qualitatively and quantitatively in this paper. Furthermore, the presence of systematic deposits of resilin in the alloscutum, which are significant for the flexibility, is demonstrated.     

Carapace repair in the green crab,Carcinus maenas (L.)

Formation of a circular hole 8–10 mm in diameter in the calcified layers of the carapace from crabs in stage C₄ of the molt cycle stimulates the tissue under and adjacent to the injury to deposit a unique calcified cuticular material below the intact membranous layer. Deposition was followed for 69 days using light microscopic histology, histochemistry, and scanning electron microscopy. Quantitative analyses of CaCO₃ were conducted using atomic absorption spectrophotometry and Gran titration. Spatial distribution of CaCO₃ was determined with X-radiography. A scab is formed by day two under the injury. At four days the epithelium changes from squamous to columnar and deposits a PAS-positive layer with an irregular lamellar fine structure, followed by highly organized lamellae structurally similar to normal exocuticle. Histochemically, however, these lamellae resemble normal endocuticle. CaCO₃ is evident external to the outermost lamellae by day eleven as a fused mass of aragonite granules. The lamellar region calcifies proximally from the outer surface and is amorphous CaCO₃. Repair cuticle is approximately 20%CaCO₃ by weight.     

Electron microscopical localization of chitin in the cuticle of Halicryptus spinulosus and Priapulus caudatus (Priapulida) using gold-labelled wheat germ agglutinin: phylogenetic implications for the evolution of the cuticle within the Nemathelminthes

 The ultrastructure of the cuticle of adult and larval Priapulus caudatus and Halicryptus spinulosus is investigated and new features of cuticle formation during moulting are described. For the localization of chitin by TEM wheat germ agglutinin coupled to colloidal gold was used as a marker. Proteinaceous layers of the cuticle are revealed by digestion with pronase. The cuticle of larval and adult specimens of both species consists of three main layers: the outer, very thin, electron-dense epicuticle, the electron-dense exocuticle and the fibrillar, electron-lucent endocuticle. Depending on the body region, the exocuticle comprises two or three sublayers. The endocuticle can be subdivided into two sublayers as well. In strengthened parts such as the teeth, the endocuticle becomes sclerotized and appears electron-dense. Only all endocuticular layers show an intense labelling with wheat germ agglutinin-gold conjugates in all investigated specimens. Additional weak labelling is observed in the exocuticle III layer of the larval lorica of P. caudatus. All other cuticular layers remain unlabelled. Chitinase dissolves the unsclerotized endocuticular layers almost completely, but also exocuticle II and partly the loricate exocuticle III. The epicuticle, the homogeneous exocuticle I and the sclerotized endocuticle are not affected by chitinase. The labelling is completely prevented in all layers after incubation with chitinase. Pronase dissolves all exocuticular layers, but not evenly. The presumably sclerotized regions of exocuticle I are not affected as well as the complete epicuticle and the endocuticle. All cuticular features of the Priapulida are compared with the cuticle of each high-ranked taxon within the Nemathelminthes with special regard to the occurrence of chitin. Based on this out-group comparison it can be concluded that: (1) a two-layered cuticle with a trilaminate epicuticle and a proteinaceous basal layer represents an autapomorphic feature of the Nemathelminthes, (2) the stem species of the Cycloneuralia have already evolved an additional basal chitinous layer, (3) such a three-layered cuticle is maintained as a plesiomophy in the ground pattern of the Scalidophora and (4) in the Nematoida, the chitinous basal layer is replaced by a collagenous one at least in the adults; the synthesis of chitin is restricted to early developmental phases or the pharyngeal cuticle. Accepted: 12 March 1998     

Structure of the integument in Paranthessius anemoniae claus,a copepod associate of the snakelocks anemone Anemonia sulcata (Pennant)

The integument of Paranthessius anemoniae has been studied with light and electron microscopy. A cuticle with clearly defined epicuticular, exocuticular and endocuticular regions overlies a cellular hypodermal layer. The distribution of carbohydrate, lipid and protein components of the cuticle were demonstrated histochemically. Parabolic striations in oblique sections of cuticle suggest that its molecular architecture fits a “twisted sheet” theory proposed for other species. Arthrodial membranes at body and limb joints have a homogeneous structure, lacking exocuticle and endocuticle. Subcuticular glands appear to secrete substances thought to be responsible for the immunity which Paranthessius seems to have to the nematocysts of its host. Small hairs, situated in cuticular cups which occur over the dorsal body surface are considered to function as rheoreceptors.     

The cytoplasmic structure hypothesis for ribosome assembly,vertical inheritance,and phylogeny

Fundamental questions in evolution concern deep divisions in the living world and vertical versus horizontal information transfer. Two contrasting views are: (i) three superkingdoms Archaea, Eubacteria, and Eukarya based on vertical inheritance of genes encoding ribosomes; versus (ii) a prokaryotic/eukaryotic dichotomy with unconstrained horizontal gene transfer (HGT) among prokaryotes. Vertical inheritance implies continuity of cytoplasmic and structural information whereas HGT transfers only DNA. By hypothesis, HGT of the translation machinery is constrained by interaction between new ribosomal gene products and vertically inherited cytoplasmic structure made largely of preexisting ribosomes. Ribosomes differentially enhance the assembly of new ribosomes made from closely related genes and inhibit the assembly of products from more distal genes. This hypothesis suggests experiments for synthetic biology: the ability of synthetic genomes to “boot,” i.e., establish hereditary continuity, will be constrained by the phylogenetic closeness of the cell “body” into which genomes are placed.     

A proposed solution to a fine-structural puzzle: The organization of gill cuticle in a crayfish (Panulirus)

Crayfish gill cuticle is approximately 2 μm thick and comprises an epicuticle and an endocuticle, which is subdivided into outer and inner layers. Sections demonstrate indistinct lamellae in the outer endocuticle and vertically striated lamellae in the inner endocuticle. Microfibrils cannot be seen in sections. Difficulties in interpretation of the fibrous architecture of the cuticle from thin sections have been overcome by examining tilted series of micrographs of sections and also by making freeze-fracture replicas of the cuticle, which reveal the microfibrils clearly. A model for the endocuticle based on a helicoidal configuration of microfibrillar laminae is proposed and the vertically striated structures seen in sections of the outer layer are accounted for by including regular rows of particles oriented perpendicular to microfibrils. The model is compared with cuticles and coverings reported from other invertebrates.     

La formation des canaux cuticulaires chez l'adulte de Tenebrio molitor L. étude ultrastructurale et remarques histochimiques

The sclerotized cuticle of adult Tenebrio shows (1) an exocuticle composed of rotating lamellate layers and of columns of cuticular material, the fibres of which run perpendicularly through the lamellae, (2) an endocuticle composed of layers with preferred orientation. In the exocuticle, the pore canals are numerous and run along the columns; they do not rotate with the lamellate layers. They show several filaments some of which leave the canals and form a dense intracuticular network. In the last layers of exocuticle, the pericolumnar canals fuse and form large endocuticular canals which rotate in phase with the cuticular fibres. The formation of columns and canals is in relation with cellular expansions which penetrate into the cuticle during cuticle deposition. Exocuticular columns seem characteristic of highly sclerotized cuticles and the intracuticular filaments may have a role in the transport of sclerotisation precursors.     

Exoskeletal structure in the Ordovician trilobite Flexicalymene

Scanning Electron Microscope study of the exoskeletal ultrastructure of secondarily phosphatized material of Flexicalymene sp. from the Upper Ordovician Maquoketa Shale, Iowa, USA, shows that the exocuticle, comprising 20% of the total exoskeletal thickness, is composed of horizontal laminar units between 0.2 and 1 μm thick. These units consist of primarily mineralized organic fibres which form horizontal laminae interconnected by inter-laminae. The endocuticle is considerably more mineralized than the exocuticle, and its original organic structure cannot be observed in untreated preparations. Etching with chromium sulphate reveals: (1) horizontal organic laminar units, 0.2 to 2 μm thick, and (2) pore canals with non-twisted walls about 0.3 μm in diameter. Exuvia cannot be distinguished from the exoskeletons of dead animals. The exoskeletal ultrastructure in trilobites agrees essentially with that in crustaceans.     

Structure of the cuticle of the common house cricket with reference to the location of lipids

Cuticle segments from the thorax, abdomen, and jumping legs of the house cricket. Acheta domesticus, were examined using histological techniques for light microscopy, scanning and transmission electron microscopy, and direct examination of frozen-fractured cuticle. The surface of untreated cuticle is covered by a lipid film which obscures fine surface detail. Standard EM preparative procedures, as well as washing the cuticle with ethanol before examination, remove this film exposing previously covered openings to dermal gland ducts and wax canals. An epicuticle, exocuticle, mesocuticle, endocuticle, and a deposition layer were present in all transverse sections of cuticle. Light microscopy showed that the exocuticle and mesocuticle are heavily impregnated with lipids, whereas there is little lipid associated with the endocuticle. Frozen-fractured cuticle clearly shows the ‘plywood’ structure of the meso- and endocuticle, while the exocuticle fractures as if it were a solid sheet. The epicuticle is composed of a dense homogeneous layer, cuticulin, outer epicuticle, and the outer membrane. Superficial wax was detected only in cuticle samples prepared using vinylcyclohexane dioxide as a polar dehydrant. The results were used to construct a comprehensive model of the cuticle of A. domesticus.     

Modified pore canals in the cuticle of Gammarus (Crustacea : Amphipoda); A study by scanning and transmission electron microscopy

A novel type of pore canal is described from the cuticle of three species of Gammarus. Each canal passes from the epidermis vertically through the endocuticle and exocuticle, and in the most distal layers of the latter is slightly expanded. Before entering the epicuticle the canal narrows, forming a neck the base of which is encircled by an electron-dense collar. Several tubular structures arise from the collar and pass distally into the reticular innermost regions of the epicuticle. Within the neck and just below its opening at the cuticle surface, a rod-like structure is inserted; this protrudes a short distance from the pore. Each pore canal is connected to many necks; the openings of the latter are aligned in rows over the surface, the openings and rows being about 0.15 and 1.0 μm apart, respectively. Changes in the pore and canal contents are visible and their significance is discussed.     

淡水育珠蚌外套膜表皮细胞分泌方式的研究

用光学显微镜、相差显微镜、透射电镜、扫描电镜对３种淡水育珠蚌外套膜表皮细胞的分泌方式进行了观察研究。几种显微镜观察对比研究的结果表明,外套膜细胞的分泌方式主要有小泡式的局部分泌;液流式、缺口式的顶浆分泌和巨浆分泌;分泌物排出时连同整个细胞一起排出和形成复层表皮后外层脱落的全浆分泌。内表皮细胞以小泡式和缺口式分泌为主;外表皮细胞多为液流式分泌和全浆分泌。内表皮分泌活动表现为持续性和连续性的特点,观察相邻细胞同时可见到各种不同的分泌时相;外表皮分泌活动呈节律性和区段性的特点,分泌活动旺盛的细胞区段和无分泌活动的区段相间出现。以上结果表明内、外表皮细胞在分泌方式及分泌物性质上均有显著差异,这种差异反映了内、外表皮功能上的不同。     

Internal anatomy of Hypochthonius rufulus (acari: Oribatida)

The internal anatomy of juveniles and adults of Hypochthonius rufulus selected as a model species representing the lower Oribatida was investigated histologically and compared with the published characteristics of higher oribatid internal anatomy. In this species, the cuticle is weak and flexible, consisting of epicuticle and endocuticle on the body, but including an exocuticle between the epicuticle and endocuticle of the legs. Walls of the mesenteron in the digestive tract are of uniform thickness and structure without any regional thickening, and there are no proventricular glands. The hindgut is apparently divided into five parts: colon 1 and 2, rectum 1 and 2, and anal atrium; food bolus exhibits a multilamellar structure in this section. The glandular system is less diversified than in some other oribatids. Tracheae are apparently lacking. Females possess only two relatively large eggs, filling one-half of opistosoma, and they lack ovipositors. Eggs are present in females during the whole year. Gonad buds appear first in the protonymph stage. Only one male was found among 146 adults studied. No male external organ (aedeagus or penis) is present.