An integumentary gland secreting a territorial marking pheromone in Atta sp.: Detailed structure and histochemistry

An abdominal pheromone-producing gland in Atta sp. was examined using light and electron microscopy techniques. The gland is composed of a bunch of juxtaposed secretory units in which the secretory ductules open on to a cribellum close to the sting base.The structure and cycles of the secreting units are described. Each includes a secretory cell with an ‘end apparatus’, ductule cells and epidermal cells. The secretory cycle of glycoproteins accumulated in the ‘end apparatus’ is discussed and a functional interpretation of the morphological components of the application system is proposed.     

Ultrastructural investigation of a salivary gland in a centipede: structure and origin of the maxilla I-gland of Scutigera coleoptrata (Chilopoda, Notostigmophora)

The maxilla I-gland of Scutigera coleoptrata was investigated using light and electron microscopy methods. This is the first ultrastructural investigation of a salivary gland in Chilopoda. The paired gland opens via the hypopharynx into the foregut and extends up to the third trunk segment. The gland is of irregular shape and consists of numerous acini consisting of several gland units. The secretion is released into an arborescent duct system. Each acinus consists of multiple of glandular units. The units are composed of three cell types: secretory cells, a single intermediary cell, and canal cells. The pear-shaped secretory cell is invaginated distally, forming an extracellular reservoir lined with microvilli, into which the secretion is released. The intermediary cell forms a conducting canal and connects the secretory cell with the canal cell. Proximally, the intermediary cell bears microvilli, whereas the distal part is covered with a distinct cuticle. The cuticle is a continuation of the cuticle of the canal cells. This investigation shows that the structure of the glandular units of the salivary maxilla I-gland is comparable to that of the glandular units of epidermal glands. Thus, it is likely that in Chilopoda salivary glands and epidermal glands share the same ground pattern. It is likely that in compound acinar glands a multiplication of secretory and duct cells has taken place, whereas the number of intermediary cells remains constant. The increase in the number of salivary acini leads to a shifting of the secretory elements away from the epidermis, deep into the head. Comparative investigations of the different head glands provide important characters for the reconstruction of myriapod phylogeny and the relationships of Myriapoda and Hexapoda.     

Clasper gland morphology and development in Potamotrygon magdalenae (Elasmobranchii: Potamotrygonidae)

Clasper gland morphology and development in Potamotrygon magdalenae and its relation with the acquisition of reproductive maturity is described in males of different developmental stages (embryos, neonates, juveniles, and reproductively active and resting adults). The glands are subcutaneous masses in the proximal base of each clasper. They are partially bilobate organs with a ventral groove that bears a row of papillae. Glands tend to be asymmetric, the left gland has a larger size, a trend that has been observed in other organs of elasmobranchs. Glands are formed by radially organized tubular secretory units lined with a simple columnar epithelium with basal nuclei and granular eosinophilic cytoplasm; vascularized loose connective tissue surrounds the gland units. The gland is covered by two layers of striated muscle tissue in circular and longitudinal arrangement. The clasper glands begin to develop in neonates and their secretory activity begins in juveniles. The active secretion of the clasper gland is observed in mature males, it includes glycoproteins and sulfated mucopolysaccharides. The size of the glands has a positive and direct relationship with body size, measured as disc width. Significant differences in clasper gland size were found between mature (active and resting) and immature (neonates and juveniles) males, suggesting that the acquisition of the sexual maturity involves the increase in the size of the gland due to a highly augmented secretory activity. Therefore, clasper glands are clearly associated with the reproductive activity of males and their secretion should have an endocrine control as other sexual secondary organs. J. Morphol. 278:369–379, 2017. © 2017 Wiley Periodicals, Inc.     

The fine structure of the colleterial glands of Hyalophora cecropia (Lepidoptera)

The structure of cells in the colleterial glands of the Cecropia silkmoth was examined. Morphologically and functionally the gland is divided into two regions, a tubular one in which columnar protein-synthesizing cells are located, and an expanded region in which flattened cells with very different structure are most prominent. The fine structure of the latter cells which are presumed to secrete a phenolic glucoside, closely resembles that of cells described in the colleterial glands of orthopterans. The protein-secreting cells have many features normally associated with pancreatic acinar, and other cells of similar function. Among these are extensive rough endoplasmic reticulum, an elaborate Golgi complex, and a modest number of mitochondria. Other features which are less usual in cells of this type are an elaborate secretory apparatus consisting of a cuticular tubule inserted into a microvilli-lined cavity at the apical end of the cell, and large numbers of cytolysomes, myelin figures, and lipid droplets. A chitogenous cell with a very distinct and specific type of ultrastructure is found associated with the secretory cell. This cell type is attached to the cuticular elements of the gland, and the main features of its cytoplasm are extensive bundles of microtubules which presumably serve as supportive elements for the secretory cells.     

Development of secretory units of mouse submandibular gland

Summary The fine structure of the secretory units of the mouse submandibular gland was studied according to the developmental sequence. The embryonic submandibular gland consists of terminal tubules and ducts. Myoepithelium is associated only with the terminal tubules, and the cells of the primary intercalated ducts show characteristics of the young striated duct cells. The major changes shortly after birth consist of: 1) opening of the secretory lumina, 2) increasing rough ER and its altered configuration, 3) dilatation of Golgi cisternae and 4) changes in the granular structure. These findings suggest that the salivary secretion first occurs after birth, and acinar differentiation or transformation of the secretory cells of the terminal tubules is induced and profoundly affected by the commencement of the secretory activity. In the intercalated ducts this process is somehow inhibited, and the granular cells found in the adult can be considered as the remnants of the secretory cells of the terminal tubules.     

Structure of the female genital glands of the oviduct in the opisthobranch mollusc, Runcina

Runcina is a small hermaphroditic opisthobranch which possesses a monaulic reproductive system. In previous studies the male copulatory apparatus, the structure of the spermatophore and also the process of oogenesis have been described. The present paper gives an account of the ultrastructure of the female genital glands of the oviduct. In Runcina the oviduct comprises three primary regions, the albumen gland, the egg capsule gland and the mucous gland. Eggs enter the fertilization chamber and as they pass the opening of the albumen gland they become surrounded by albumen or perivitelline fluid. The eggs appear to become encapsulated as they traverse the egg-capsule gland and are eventually stuck together by mucus to form an egg mass. The epithelial lining of the three glands consists of alternating ciliated and secretory cells. The characteristics in secretory products of the glandular cells are described, and are discussed with reference to the way they contribute to egg vestment.     

The fine structure of the vesicular component of the ultimobranchial gland of the domestic fowl

Summary The ultrastructure of the polymorphic vesicular component of the ultimobranchial gland of the domestic fowl (Gallus gallus domesticus) has been described in detail, together with the structure of the cell strands interconnecting the vesicles and the parathyroid nodules lying within the ultimobranchial stroma. The vesicles frequently appear to arise from the nodules by way of the cell strands. The strands show a structure of their component cells intermediate between that of the parathyroid and the vesicular cells, although the position at which the strand changes from an essentially parathyroid structure to an essentially vesicular structure is very variable. The degree and kind of secretory activity within different cell types has been described. A review of the structure of ultimobranchial glands throughout the vertebrates shows that similar tissue with a similar secretory potential has been observed in all vertebrate classes, suggesting a functional significance for this part of the gland.     

Morphology and ultrastructure of a secretory region enclosed by the venom reservoir in social wasps (Insecta,Hymenoptera)

The morphology and ultrastructure of the convoluted gland inside the venom reservoir of four species of social Vespidae are described. The cells of the venom gland (including the convoluted gland) can be divided into six groups: (1) epithelial cells, (2) glandular cells with the end apparatus secreting into the tubule inside the convoluted gland (internal or embedded tubule), (3) a continuous arrangement of glandular cells with the end apparatus secreting directly into the venom reservoir, (4) glandular cells that are loosely dispersed along the tubule lumen between the free tubules and the embedded tubule of the convoluted gland, (5) secretory cells of the free tubules and (6) duct cells. One kind of secretory cell, hitherto unknown and described in this paper (group 3), is characterized by the presence of a well-developed end apparatus, usually with enlarged extracellular spaces, but lacking the normally associated duct cells. The secretory cells contain several stacks of granular endoplasmic reticulum, but these are mainly concentrated in the middle of the cell. The basal half of the cells contains many lipid droplets. Although the function of the convoluted gland is not yet understood, an hypothesis is related to what is known of the function of reservoir secretory cells in solitary wasps. All wasp species studied showed the same organization of the convoluted gland, which clearly distinguishes their venom gland from that of Sphecidae.     

Morphology,histochemistry and physiology of the major salivary glands in the echidna Tachyglossus aculeatus (Monotremata)

The three major salivary glands of the monotreme echidna are described. The parotid is a typical serous gland with tubulo-acinar secretory endpieces and a well-developed system of striated ducts. The mandibular gland, although light microscopically resembling a mucous gland, secretes very little glycoprotein. Its cells are packed instead with serous granules, resembling in fine structure the “bull's eye” granules in the mandibular gland of the European hedgehog Erinaceus europaeus. The sublingual glands secrete an extremely viscous mucous saliva. Expulsion of this saliva through the narrow ducts is probably aided by contraction of the extensive myoepithelial sheaths surrounding the secretory tubules. Application of the glyoxylic acid induced fluorescence method failed to demonstrate adrenergic innervation in any of the glands.     

Structure of the smallest salivary-gland secretory protein gene in Chironomus tentans

The salivary gland secretion in the dipteran Chironomus tentans is composed of approximately 15 different secretory proteins. The most well known of the corresponding genes are the four closely related Balbiani ring (BR) genes, in which the main part of each approximately 40-kb gene is composed of tandemly arranged repetitive units. Six of the seven additional secretory protein genes described share structural similarities with the BR genes and are members of the same BR multigene family. Here we report the identification of a new secretory protein gene, the spl2 gene, encoding the smallest component of the C. tentans salivary gland secretion. The gene has a corresponding mRNA length of approximately 0.7 kb and codes for a protein with a calculated molecular weight of 7,619 Da. The sp12 gene was characterized in seven Chironomus species. Based on a comparison of the orthologous gene sequences, we conclude that the sp12 gene has a repetitive structure consisting of diverged 21-by-long repeats. The repeat structure and the codon composition are similar to the so-called SR regions of the BR genes and the sp 12 gene may represent a diverged member of the BR multigene family. Correspondence to: L. Wieslander     

Ultrastructure of epidermal glands in neotenic reproductives of the termite Prorhinotermes simplex (Isoptera: Rhinotermitidae)

The ultrastructure of epidermal glands in neotenic reproductives of Prorhinotermes simplex is described and their development is compared among young and old neotenics of both sexes. Secretory cells forming the epidermal gland are attached to the cuticle all over the body. The glands are formed by class 1 and class 3 secretory cells and corresponding canal cells with secretory function. Class 1 cells are sandglass-like and class 3 secretory units are located among them. Class 1 cells contain predominantly tubular endoplasmic reticulum, the major part represents the smooth and the minor the rough form. Numerous electron dense granules occur in the cytoplasm, they are always disintegrated prior to be released. Class 3 secretory cells contain a large amount of vacuoles, which are always lucent in males while newly produced vacuoles are dense in females. Dense vacuoles are frequently transformed into lucent ones before being released. Canal cells are locally equipped with microvilli. The conducting canal is surrounded by an electron dense secretion of regular inner structure. The cytoplasm of the canal cell contains numerous mitochondria, rough endoplasmic reticulum and a large proportion of microtubules. The young neotenic reproductives differ from the old ones by a lower amount of secretory products. Epidermal glands probably produce substances inhibiting the occurrence of superfluous reproductives.     

The ultrastructure of the female accessory gland,the cement gland,in the insect Rhodnius prolixus

The cement gland of Rhodnius prolixus is an epidermally derived tubular gland consisting of a distal synthetic region and a proximal muscular duct region. The synthetic region consists of numerous secretory units joined to a central chitinous duct via cuticular ductules. Proteinaceous secretion, synthesized by the goblet-shaped secretory cell, passes through the delicate cuticular lattice of a ductule-end apparatus and out through fine ductules to the central duct. Secretory cells are rich in rough endoplasmic reticulum and mitochondria. Light microscopy, SEM and TEM reveal the delicate lattice-like end apparatus structure, its formation and relationship to the secretory cell. The secretory cell associates via septate junctions with a tubular ductule cell that encloses a cuticle-lined ductule by forming an elaborate septate junction with itself. The ductules are continuous with the cuticle lining of the large central duct that conveys secretion to the proximal area. The proximal muscular duct has a corrugated cuticular lining, a thin epithelium rich in microtubules and thick longitudinal, striated muscles which contract during oviposition, forcing the secretion out. Histochemistry and electrophoresis reveal the secretion as proteinaceous.     

The secretory cycle of a gland involved in pheromone production in the noctuid moth, Pseudaletia separata

The structure of the pheromone producing gland is briefly described. Examination of variations in gland volume, nucleus to cytoplasm ratio, and distribution and form of nucleic acids suggests that the secretory cycle of this gland consists of three phases. Two types of small vesicle are found and suggested as possible sites for pheromone metabolism.     

Structural and functional aspects of salivary fluid section in Calliphora

Calliphora salivary glands are described, emphasizing correlations between structure and physiology. In vitro studies show that the distal part of each gland produces a potassium-rich primary saliva when stimulated with 5-hydroxytryptamine or cyclic AMP. The secretory cells have elaborate canaliculi opening into the lumen. Stimulation of the secretory region causes a 60-fold increase in fluid secretion rate without affecting cell structure. The proximal part of the gland reabsorbs potassium when stimulated with cyclic AMP, but 5-HT has no effect. Potassium reabsorption from the primary saliva results in formation of a dilute saliva. The structure of the secretory and reabsorptive cells is discussed with regard to the functional role of long narrow channels in transport.     

Distribution and comparative morphology of the cloacal gland in ants (Hymenoptera : Formicidae)

The cloacal gland is a paired exocrine structure, which has so far been described only in the formicine species, Camponotus ephippium and Cataglyphis savignyi (Hymenoptera : Formicidae). The gland is formed by 2 clusters of bicellular units with slender duct cells, releasing the glandular secretion through the cloacal membrane. In the present work, a number of ant species, largely of the Formicinae subfamily, have been surveyed for the presence of a cloacal gland. The gland is present in nearly all formicines screened, albeit with a variable development. Cataglyphis, one of the genera with a very prominent cloacal gland, was chosen for a more detailed comparative study. At the ultrastructural level, secretory cells were observed having a well-developed smooth endoplasmic reticulum and Golgi apparatus, typical for pheromone-producing glandular cells. The gland is also present in all dolichoderines screened, but in none of the species of the Aneuretinae, Myrmeciinae, Myrmicinae, Nothomyrmeciinae, or Pseudomyrmecinae investigated. This provides tentative evidence that the cloacal gland is a synapomorphy of the Formicinae and Dolichoderinae, giving support for their hypothesized sister group relationship. Up to now, the function of the cloacal gland remains largely enigmatic.     

Seasonal changes in the structure and secretory activity of the androgenic gland of Travancoriana schirnerae

This study investigated the seasonal variation in the structure and secretory activity of the androgenic gland (AG) in the freshwater crab: Travancoriana schirnerae. The androgenic gland is an elongate structure, attached to one side on the wall of the ejaculatory duct. Histological studies showed the presence of three cell types, which differ in size, shape of nuclei, and presence or absence of secretory vesicles. Type I cells are small with large nuclei whereas type II cells are large with small nuclei. Type III cells are intermediate in size and exhibited streak-like nuclei and transparent cytoplasm. Seasonal changes were discerned in the morphology, histology and secretory activity of the gland. March-June appeared to be the active season with type II cells containing secretory vesicles. The mode of release of secretion found to be holocrine. The secretory activity almost completed by July-August (the mating season) with vacuolization of type II cells. The gland remained inactive from September-December with abundance of vacuoles, scattered pycnotic nuclei, indistinct cell membranes and total cellular degeneration. January-February was the revival period with type I cell proliferation. The present study revealed that the secretory activity of the gland is in tune with the male reproductive cycle.     

The comparative morphology of epidermal glands in Pentatomoidea (Heteroptera)

The Heteroptera show a diversity of glands associated with the epidermis. They have multiple roles including the production of noxious scents. Here, we examine the cellular arrangement and cytoskeletal components of the scent glands of pentatomoid Heteroptera in three families, Pentatomidae (stink bugs), Tessaratomidae, and Scutelleridae (shield-backed bugs or jewel bugs). The glands are; (1) the dorsal abdominal glands, (2) the tubular glands of the composite metathoracic gland, and (3) the accessory gland component of the composite metathoracic gland. The dorsal abdominal glands are at their largest in nymphs and decrease in size in adults. The metathoracic gland is an adult-specific gland unit with a reservoir and multiple types of gland cells. The accessory gland is composed of many unicellular glands concentrated in a sinuous line across the reservoir wall. The lateral tubular gland is composed of two-cell units. The dorsal abdominal glands of nymphs are composed of three-cell units with a prominent cuticular component derived from the saccule cell sitting between the duct and receiving canal. The cuticular components that channel secretion from the microvilli of the secretory cell to the exterior differ in the three gland types. The significance of the numbers of cells comprising gland units is related to the role of cells in regenerating the cuticular components of the glands at moulting in nymphs.     

The subepithelial gland in ants: a novel exocrine gland closely associated with the cuticle surface

Two glandular systems were discovered that secrete their products onto the cuticular surface in ants. The first, the subepithelial gland, was previously undescribed in ants, and is found throughout the body just beneath the epithelium. This gland consists of independent secretory units, each made up of a single gland cell and an associated duct cell that penetrates the cuticle. Its ultrastructural appearance is consistent with possible hydrocarbon production. Examining 84 ant species, the subepithelial gland was found in eight subfamilies (out of 13), although not necessarily in all species. In a single ant species, Harpegnathos saltator, it was the epithelium itself that was enlarged and functioned as a gland. The enlarged epithelial cells secrete their products directly onto the cuticle through distinct cuticular crevasses.     

Fine Structure of the Esophagus of Pratylenchus penetrans

The fine structure of the esophagus of Pratylenchus penetrans is described. The gland lobe is syncytial and contains two types of nuclei: three large nuclei with little chromatin, and more numerous smaller nuclei with large amounts of chromatin. Some of the smaller nuclei are associated only with glandular tissue, whereas others are part of nerve ceils within the esophagus. Clusters of free ribosomes, rough endoplasmic reticulum, and numerous mitochondria occur in the lobe region where the secretory granules are formed. No Golgi bodies were observed. On the basis of these observations, possible differences in the mechanism of secretory granule formation between plant-parasitic nematodes are discussed. Several other minor differences between the fine structure of other plant-parasitic nematodes previously examined and that of P. penetrans are also noted.     

Morphology of the aedeagal gland of the male mealworm beetle (Tenebrio molitor L.)

The aedeagal gland of male Tenebrio molitor consists of numerous acini containing several secretory units (organules) of three epithelial cells in series. The distal cortical cell and intermediate cell are secretory cells. Secretory products are passed into microvilli-lined extracellular reservoirs. From these storage areas products flow through minute canaliculi and into the efferent ductule. Canaliculi, cuticular trabeculae, and fibrillar material are characteristic features of the efferent ductules within the extracellular reservoirs of secretory cells. After passing from the secretory cells, the efferent ductule penetrates the basal ductule cell. The thin epicuticle that comprises the wall of the ductule is confluent with the epicuticle of the cuticular sheath forming the wall of the genital pocket. Secretory products flow from the cortical cell ductule into the intermediate cell and eventually empty into the genital pocket. A chemical reaction apparently takes place in the intermediate cell ductule, resulting in a frothy secretion product. When released from the ductule, this frothy product forms a foam-like layer that coats the inner wall of the genital pocket. Ultrastructural and probable functional aspects of this gland are described and discussed.