An ultrastructural study of the larval integument of the midge,Chironomus riparius meigen (Diptera: Chironomidae)

Summary The larval integument of the midge, Chironomus riparius Mg., is unusually thin although it conforms with the normal insect pattern. The cuticle of the post-cephalic segments is about 3 m thick and overlies an epidermis which has an irregular basal plasma membrane resulting in spaces occurring between it and the basement membrane. The ventral tubuli have a similar epidermis but the cuticle is somewhat thinner. The anal papillae have the thinnest cuticular covering with a uniquely folded epicuticle of variable thickness, and their epidermis has the characteristics of a transporting epithelium. No evidence of pore canals could be found in the cuticle of any part except the head capsule which has a remarkably smooth epicuticle and a distinct layer which may represent the exocuticle. There are no spaces between the basement membrane and basal plasma membrane of the epidermis in the head. Ultrastructural evidence would suggest that gaseous exchange can occur across most of the post-cephalic integument.The author is indebted to Mrs. L. Rolph and Mr. R.L. Jones for their technical assistance     

The structure of the integument of the sea cucumber,Thyone briareus

The integument and podia of the sea cucumber Thyone briareus were examined by bright field and electron microscopy. The epidermal surface was found to be covered by an acellular, PAS positive cuticle which appeared to be secreted by the underlying epidermal cells. Although the superficial portion of the cuticle contains numerous fine filaments, their ultrastructure bears no resemblance to collagen fibers. The epidermal cells are widely spaced and have long apical processes that extend along the under surface of the cuticle forming a contiguous epithelium. The apical expansions of the epidermal cells are attached to one another by means of septate desmosomes which may run continuously around all epidermal cells. Special attachment structures within these apical expansions appear to bind the cuticle to the dermis. The epidermal cells and their apical expansions are separated from the dermis by an 800 Å thick basement membrane. Granule containing cells in the upper dermis send processes up to the cuticle where they are bound to the typical epidermal cells by septate desmosomes. The abundant membrane bound granules of the cells enter villous-like processes which pass through the cuticle. The function of these cells may be to produce an adhesive material on the podia or they may be pigment cells. The thick dermis consists of a superficial zone, containing largely ground substance; a middle or laminated zone containing laminae of collagen fibers arranged in an orthogonal fashion; and a hypodermis consisting largely of ground substance and reticular fibers. Fibroblasts are abundant in the superficial dermis and between the collagen laminae. Wandering coelomocytes, or morula cells, accumulate between the collagen laminae and in the hypodermis. They may also become an integral part of the epidermis by forming septate desmosomes with epidermal cells. Morula cells contain highly specialized spherules whose tinctorial properties and electron microscopic appearance suggest that they contain protein and mucopolysaccharide.     

Cuticular encystment of Autographa nigrisigna eggs by epidermal cell migration

It was previously established that Autographa nigrisigna loopers form cuticular cysts at the dorsal site of the 9th (penultimate) abdominal segment after parasitization by the solitary endoparasitoid Campoletis chlorideae and get rid of the parasitoid egg with the old cuticle at ecdysis. The cuticular cyst consists of a space between the old cuticle and new cuticle formed by the epidermis to enclose the parasitoid egg. The fact that A. nigrisigna loopers exclude the oviposited egg from the hemocoel using a cuticular cyst raises the question how the parasitoid egg passes through the epidermis. To exclude the endoparasitoid eggs from the hemocoel, the epidermis is required to move the location of the parasitoid egg. In the current study, we investigated the morphological process of cuticular cyst formation. First, the oviposited egg drifted to the 9th abdominal segment located at the open end of the dorsal vessel as a result of force generated by the hemolymph current from the oviposition site, and formed contacts with the integument containing the fat body (FB). The epidermis, in contact with the egg, then began to move along with the basement membrane formed on the surface of the FB, and settled under the egg, thus altering its location. This inversion was duplicated in vitro using integument from the 9th abdominal segment when parasitoid eggs were inserted between the epidermis and FB. When the integument, without the FB, was incubated on an agar plate, the epidermal cells migrated on the plate. Integument without eggs showed no signs of migration from their original sites. When the actin polymerization inhibitor latrunculin B was added to the cultures, the epidermal cells remained in their original location.     

The structure of developing muscle insertions in insects

The relationship between muscles and the components of the integument in muscle insertions have been studied with the electron microscope in two insects, Calpodes ethlius (Hesperiidae, Lepidoptera) and Rhodnius prolixus (Reduviidae, Hemiptera). The area of contact between the muscles and the epidermis is increased by interdigitating processes whose membranes are joined by intermediate junctions. The junctions occur at the level of a Z line so that actin filaments attach directly to them. Within the epidermis, microtubules extend from the junctions of the myoepidermal connection to the cuticle, where they attach to hemidesmosomes which line deep indentations of the membrane. The microtubules probably enable the tendinous epidermal cells to withstand the tensions exerted upon them by the muscles. The epidermis is anchored to the cuticle by tonofibrillae, homogeneous rods secreted in the deep indentations of the plasma membrane. Since the tonofibrillae of successive instars are continous, they penetrate and attach to the cuticulin, the outermost layer of the epicuticle.     

Integument of Kamptozoa Barentsia discreta Busk, 1886

The integument of the colonial species Barentsia discreta has been investigated in the present work. On its greater length the integument is presented by a monolayered unciliated epithelium covered by a layer of microvillar cuticle. The floor of the atrial cavity and the frontal surface of tentacles is lined by ciliated epidermis covered by a protocuticle. Sensitive and secretory cells are present in the epidermis.     

An ultrastructural analysis of the cuticle,epidermis and esophageal epithelium of Eisenia foetida (Oligochaeta)

The epidermis of Eisenia is covered by a cuticle and rests on a basement lamella. The cuticle, which is resistant to a variety of enzymes, is composed of non-striated, bundles of probable collagen fibers that are orthogonally oriented and are embedded in a proteoglycan matrix. The basement lamella consists of striated collagen fibers with a 560 Å major periodicity. Proximity and morphology suggest that the epidermis may contribute to both the cuticle and the basement lamella — that is, the single tissue may synthesize at least two types of collagen. The epidermis is a pseudostratified epithelium containing three major cell types (columnar, basal and gland) and a rare fourth type with apical cilia. The esophagus is lined by a simple cuticulated epithelium composed predominantly of a single cell type, which resembles the epidermal columnar cell. Rare gland cells occur in the esophageal epithelium, but basal cells are lacking.     

The fine structure of epidermal papillae of Travisia forbesii (Annelida)

The structure of the epidermis of Travisia forbesii was described using light and electron microscopy. The epidermis is a highly modified variant of the normal one-layer polychaete epithelium. It consists of basal epidermal cells and an external layer of closely sited papillae consisting of glandular and supportive epidermal cells, and extensive electron-transparent intercellular spaces. The papillae are embedded in the thick cuticle. Each papilla has a peduncle, which is formed by one cell that penetrates the inner cuticle layer to the basal epidermal cells. A fold of basement membrane forms the core of the peduncle and ends in the base of a papilla. All epidermal cells are connected to each other with apical cell junctions and to the basement membrane with hemidesmosomes, so the epithelium is continuous and uninterrupted. The epidermis has an intra-epidermal neuron plexus. The structure of the papillae is compared with papillae and tubercles of other polychaetes, and the possible functional significance and phylogenetic implications of these structures are discussed.     

Zur Feinstruktur des Integumentes von Trachydemus giganteus Zelinka (Kinorhyncha)

The integument of the Kinorhynch Trachydemus giganteus was investigated by means of electron microscope. It was found that the epidermis is composed of individual cells. The cuticle and receptor cells, which in part bear an apical cilium, exhibit arthropod characteristics.

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Mit Unterstützung durch die Deutsche Forschungsgemeinschaft.     

Integument and epidermal sensory structures of Myzostoma cirriferum (Myzostomida)

Summary The fine structure of the integument of Myzostoma cirriferum is described with special attention to the integument sensory areas. Hypotheses about the function and a functional model of these are proposed. The integument consists of an external pseudostratified epithelium with cuticle (the epidermis) covering a parenchymo-muscular layer (the dermis). The dermis includes two types of cells: muscular fibers of the double obliquely striated type and parenchymal cells. Differences occur in the epidermis, which consists either of a large non-innervated myoepithelial area (viz. the regular epidermis). or of several rather localized sensory-secretory areas associated with discrete nerve proceses (viz. the sensory epidermis). The regular epidermis is made up of three types of cell: covering cells, ciliated cells and myoepithelial cells. The sensory epidermis shows small or marked structural variations from the regular epidermis. Small variations occur in the cirri, the buccal papilla, the body margin, the parapodia and the parapodial folds where nerve processes insinuate between epidermal cells. They are thought to be mechanoreceptor sites that could give information on the structural variations of the host's integument and participate in the recognition of individuals of the same species. The sensory epidermis differs markedly from the regular eidermis in the four pairs of lateral organs. Each lateral organ consists of a villous and ciliated dome-like central part, surrounded by a peripheral fold. The epidermis of the fold's inner part (viz. the part facing the central dome) is made up of secretory cells, while that of the fold's outer part is similar to the regular epidermis. The epidermis of the dome includes vacuolar cells, sensory cells and a different type of secretory cell. Lateral organs are presumed to be both chemoreceptors and mechanoreceptors. They could allow the myzostomids to recognize the host's integument and prevent them from shifting on the surrounding inhospitable substrate.     

Ultrastructural observations on the gills of polychaetes

The gills of several polychaete species belonging to 9 families were studied by scanning and transmission electron microscopy. The surface epithelium is covered by a thin cuticle which is invaded by microvilli penetrating the epicuticle in certain species. Some epithelial cells bear cilia, others are mucus-producing cells. The ciliary cells may be arranged in rows and maintain a constant flow of water over the gills. The distance between external water and blood stream differs considerably according to the species investigated. InMalacoceros the gills are characterized by closed afferent and efferent subepithelial vessels, which correspond to tubular invaginations of the coelomic wall. These vessels are lined by the basement lamina of the coelothelial cells, which are of the epitheliomuscular type. The vessels are open in the gills of other polychaetes and release the blood stream into a system of spaces immediately below the epidermis (e.g. in the branchial lamellae ofPectinaria andTerebellides). In several species the blood comes into very intimate contact with the cuticle (e.g. in the gill filaments ofDendronereides), but also in these animals both are separated by a very small epidermal layer.Supported by DFG Sto 75/3-6.     

Ultrastructure of the nymphal integument in the camel tick Hyalomma (Hyalomma) dromedarii (Ixodoidea: Ixodidae)

The ultrastructure of the nymphal integument in the ixodid tick Hyalomma (Hyalomma) dromedarii is compared for stages of development during and after feeding, and up to the first step of molting, apolysis. The integument comprises a cuticular layer and underlying epidermal cells. The body cuticle, which consists of both sclerotized and non-sclerotized parts, is divided into an outer, thin epicuticle, and an inner, thick, fibrillar procuticle. Pore canals in the procuticle are continuous with wax canals which traverse the epicuticle. As feeding progresses, the parallel, extensible epicuticular folds disappear due to the gut filling with ingested blood. The procuticular zone, however, becomes subdivided into an exocuticle, similar to the previously seen procuticle, and a lamellate endocuticle. Pore canals lose their parallel pattern and appear to have become deformed by stretching of the cuticle. The flat epidermal cells grow asynchronously during feeding; their cytoplasm becomes packed with well-developed rough endoplasmic reticulum (RER), while the cell apices project long microvilli extending deep into the procuticle. The RER undergoes ultrastructural changes indicating synthetic activity. Dense material released through the microvilli may serve to lyse the endocuticle and thus cause separation of the cuticle from the epidermis during apolysis. The lysed area, the exuvial cavity, is filled with lysed components which are probably withdrawn by endocytosis into the apical coated vesicles which appear in epidermal cells. Two types of integumental glands, which may participate in wax production, are observed in this study. The ultrastructure of their previously undescribed cuticular ducts is described, in addition to other hypodermal structures including epidermis-muscle attachments and sensory receptors.     

Ultrastructure of the epidermis in the ice worm, Mesenchytraeus solifugus

The ice worm is adapted for life at O°C. A survey of the ultrastructure of the cuticle, epidermal epithelium and basement membrane does not reveal any features which self-evidently correlate with such metabolic specialization; instead, these tissues are much like those of the earthworm and some freshwater oligochaetes. The cuticular fibers are unstriated. Epithelial cells aresuggested as the source of cuticular material. Epithelial microvilli penetrate the cuticle. There is an array of membrane bound bodies on the cuticle surface. The basement membrane fibers are transversely striated and are oriented in crossed lamellae. The junctional complex is represented by azonula adhaerens and septate desmosome.     

A radioautographic study of collagen synthesis by earthworm epidermis

Secreation by the epidermis of two oligochaetes (Eisenia and Enchytraeus) was investigated radioautographically following administration of 3H-proline, 3H-tryptophan or Na2(35)SO4. Regionally epidermal columnar cells of Enchytraeus synthesize the overlying, probably collagenous, cuticle. Eisenia epidermis does not recordably synthesize the cuticle until after wounding (first eight segments removed). By two days postoperative the epidermal columnar cells of Eisenia synthesize the collagenous cuticle and, later in regeneration, the epidermis may simultaneously synthesize the different collagen of the underlying basement lamella. The epidermis of Enchytraeus, but not of Eisenia, synthesizes some sulfated material associated with the cuticle surface.     

Why caterpillars do not grow short and fat

The integument of a caterpillar consists of a cuticle that is structurally isotropic in the plane of the surface, overlaying an epidermis and basal lamina. Between ecdyses the cuticle stretches so that the proportions of caterpillars are about the same at the end of a stadium as at the beginning. This introduces a paradox, since under uniform internal hydrostatic pressure, the hoop stress is twice the axial stress. Stress in the integument of a caterpillar inflated by compressed hemolymph would be twice as great in the circumference as in the axis, causing it to become progressively fatter in proportion to its length as it grows. This does not happen because axial pleats allow caterpillar cuticle to extend more easily in the axis, allowing uniform growth in spite of the greater hoop stress from internal hydrostatic pressure.     

Immunoelectron microscopic localization of actin in migrating cells during planarian wound healing

Wound repair in planarians is mainly characterized by two cell-migratory events involving the epidermis adjacent to the wound and its basement membrane. The first event is the migration of epidermal cells to cover the wound surface; the second one is the migration of newly differentiating replacement epidermal cells from the parenchyma to the epidermis. In addition to these events, migration of fixed parenchymal cells is observed during wound healing. All migrating cells were characterized by the presence of actin, as shown by the results obtained by means of indirect immunolocalization with fluorescent and electron microscopy. Migrating cells were heavily labeled with gold particles, which clustered at the level of cell-matrix and cell-cell contacts.     

The liquid crystalline nature of the cytoskeleton in epidermal cells of the chaetognath Sagitta

The chaetognaths have a multilayered epidermis, which is not covered by cuticle, except in the head region. Two kinds of cells are found in the epidermis: the filament-rich cells, adjacent to the basement membrane, and superficial cells, which are filament poor. The filament-rich cells, which are linked by gap junctions and columnar junctions, are highly developed in the collarette region, which joins the head and the trunk. As elsewhere in the epidermis these cells are covered by the filament poor cells which are linked by zonulae adhaerentes, gap junctions and septate junctions. The filaments present in the inner cells of the collarette form a twisted fibrous arrangement, which shows parallel series of nested arcs when observed in oblique section. Such systems are well known in numerous skeletal materials and correspond to polymerized analogues of certain liquid crystals. The amount of connective tissue is extremely reduced in Sagitta. One can hypothesize that filament-rich cells are abundant in regions which undergo strong deformations. This is the case in the collarette, in contact with the basement membrane of the epidermis (which in turn is in contact with a myotendinous system), in a region where ingested prey must go through the general cavity where there is high internal pressure.     

Etude ultrastructurale de l'évolution du tégument de la Sangsue Hirudo medicinalis L. (Annélide,Hirudinée) au cours d'un cycle de mue / Developmental changes in the integument of the leech Hirudo medicinalis L. (Annelida,Hirudinea) during a molting cycle. An ultrastructural study

Summary

The integument of the leech Hirudo medicinalis is mainly composed of a single layer of cuticle-secreting epidermal cells. The cuticle is made up of collagen fibers which support a layer of membrane-bound epicuticular projections.

Shedding of the old cuticle is preceded by the formation of a new cuticle. The epicuticular projections are the first to develop: they originate from the tips of numerous microvilli of the epidermal cells. As soon as it appears, the newly-formed collagen layer is firmly attached to the epidermal cells by numerous hemidesmo-somes, whereas the old cuticle is no longer connected with the epidermal surface. The epidermal cells exhibit marked characteristics of secretory activity during the laying down of the new cuticle.

The observations are discussed in connexion with recent findings of high ecdysteroid levels in leeches at the beginning of the molting cycle.     

Integument and hemocyte peptides

There are four routing classes of integument peptide in the caterpillar of Calpodes ethlius. The epidermis secretes peptides apically into the cuticle (C), basally into the hemolymph (H) and in both directions (BD). Peptides in a 4th class (T), are presumed to be transported across the epidermis, because the epidermis does not synthesize them although they occur in both cuticle and hemolymph. In a search for the origin of the presumed transepidermal peptides we found that hemocytes contain some peptides from all four routing classes. Peptides prepared from washed hemocytes reacted in immunoblots to antibodies against integument peptides prepared from hemolymph and cuticle. These peptides are probably synthesized by hemocytes because they matched those from medium containing [³⁵S]methionine in which hemocytes had been incubated. Calpodes hemolymph contains four hemocyte types. Immunogold labelling localized integument peptides in the secretory pathway of granulocytes and spherulocytes and in the cytosol of oenocytoids but not in plasmatocytes. Each peptide was localized in a particular kind or kinds of hemocyte. Granulocyte secretory vesicles reacted with antibodies to C180, C55 and BD82 kDa peptides. Spherulocytes secretory vesicles reacted with antibodies to C180, C55, BD89, BD82 and a 78 kDa peptide presumed to be the precursor of T66. Oenocyotoids reacted with antibodies to H45, 38, 32, 23 and BD89 kDa peptides. Spherulocytes were the only tissue to react with antibodies to the T66 kDa peptide that is found abundantly in cuticle and hemolymph. Spherulocytes are therefore presumed to secrete the 66 kDa peptide into the hemolymph from where it is transported to the cuticle. The C180 and C55 kDa peptides do not occur in hemolymph. Their presence in granulocytes and spherulocytes may be associated with hemocyte functions such as basal lamina formation, since immunogold localized them in that part of the basal lamina next to the hemolymph, as would be expected if hemocytes deposited components onto the exposed hemolymph surface. The presence of hemolymph peptides in oenocytoids is more difficult to interpret, since the antigenic reactions are localized in the cytosol rather than in the secretory pathway expected for exported proteins. We conclude that integument peptides are not secreted only by the epidermis, nor is the cuticle their only destination.     

Can larval epidermis omit the secretion of a pupal cuticle in a saturniid moth, Samia cynthia ricini?

When the larval tissue is exposed to the hormonal milieu lacking juvenile hormone, adult characters appear directly omitting the pupal stage in some insects but not in others. In Samia cynthia ricini, a species belonging to the latter group, a possible omission of pupal characters was tested by previously untried experiments. Firstly, the possibility that the larval epidermis of only some stages is capable of responding so as to omit to secrete the pupal cuticle was tested. Pieces of larval integument taken from various developmental stages were implanted into developing (pharate) adults. None of these failed to secrete the pupal cuticle. Secondly, pieces of larval integument were first implanted into brainless pupae and left there for a month to eliminate the effect of a trace of juvenile hormone which might have been carried over by the implants. They were then caused to develop, and they again secreted pupal cuticle. It is concluded that the larval epidermis cannot omit secreting pupal cuticle in this species.     

A fine structural analysis of the epidermis and cuticle of the oligochaete aeolosoma Bengalense stephenson

The fine structure of the epidermis and cuticle has been described for the oligochaete Aeolosoma bengalense. The epidermis is a pseudostratified epithelium and consists of the following cell types: ciliated and nonciliated supportive cells, pigment cells and associated satellite cells, mucous cells, basal cells, and ciliated non-supportive columnar cells. Overlying and restricted to the supportive cells is a delicate cuticle composed of: (a) a discontinuous layer of membrane-bounded surface particles; (b) a thin filamentous layer of moderate electron density just under the surface particles; (c) a thicker inner filamentous layer of low electron density. Digestion with pronase effectively removes the cuticle. This, together with the fact that it stains with alcian blue and ruthenium red, indicates that the cuticle contains an acid mucopolysaccharide. Regeneration of the cuticle, following pronase treatment, is marked by the elaboration of numerous microvilli by the supportive cells. Most of the microvilli are transitory and evidence supports a microvillar origin for the cuticular surface particles. The presence of cuticular surface particles may be a characteristic shared in common by all oligochaetes and, perhaps, some polychaetes.