An interommatidial exocrine gland with a “nail-headed” structure in the water strider Aquarius remigis (Hemiptera,Gerridae)

The fine structure of the interommatidial exocrine glands, found in the compound eyes of the water strider Aquarius remigis, is described using light, scanning, and transmission electron microscopy. The glandular pores of the glands are specialized into minute “nail-headed” structures (NS), which are described for the first time in arthropod compound eyes. Each NS is composed of two components: a rod-like stalk and a cup-like depression. The TEM study shows that the glands are class 3 epidermal glands as defined by Noirot and Quennedey (1974, 1991). Each gland consists of 3 cells: a gland cell, an intermediary cell, and a duct (canal) cell. The gland cell contains abundant electron-lucent vesicles, while the intermediary cell contains a large number of osmiophilic secretory granules. These two cells might secrete different substances which mix together in the dilated sac-like portion of the conducting canal before final release. The possible functions of the secretions released from these glands are discussed.     

The “rosette-like” structures in the cuticle of Petrobius brevistylis are the openings of epidermal glands

We examined the “rosette-like” structures (RS), found in Archaeognatha and Thysanura, in the compound eyes and the antennae of the machilid Petrobius brevistylis using SEM and TEM. The nature of the RS was unknown until now, and hypothesized to be either a sensillum or the opening of a gland. Our studies show that RS are the orifices of epidermal glands. A gland consists of a single glandular unit of 4 cells: a duct cell, a secretory cell, a ciliary cell and an enveloping cell. The glands are class 3 epidermal glands as defined by Noirot and Quennedey (1974).     

Ultrastructure of male accessory glands in the scorpionfly Sinopanorpa tincta (Navás, 1931) (Mecoptera: Panorpidae)

The ultrastructure of male reproductive accessory glands was investigated in the scorpionfly Sinopanorpa tincta (Navás, 1931) (Mecoptera: Panorpidae) using light and transmission electron microscopy. The male accessory glands comprise one pair of mesodermal glands (mesadenia) and six pairs of ectodermal glands (ectadenia). The former opens into the vasa deferentia and the latter into the ejaculatory sac. The mesadenia consist of a mono-layered elongated columnar epithelium, the cells of which are highly microvillated and extrude secretory granules by means of merocrine mechanisms. The epithelium of ectadenia consists of two types of cells: the large secretory cells and the thin duct-forming cells. These two types of cells that join with a cuticular duct constitute a functional glandular unit, corresponding to the class III glandular cell type of Noirot and Quennedey. The cuticular duct consists of a receiving canal and a conducting canal. The secretory granules were taken up by the receiving canal and then plunged into the lumen through the conducting canal.     

Morphology and ultrastructure of the abdominal integumentary glands of Acanthoscelides obtectus Say (Coleoptera : Bruchidae)

The morphology of the abdominal integumentary glands of the bean weevil Acanthoscelides obtectus (Coleoptera : Bruchidae) is described. There are 2 types: glands with long secretory ducts and ampullate glands with short ducts. The former are distributed throughout the integument and have 2 secretory cells, one of which is connected to the reservoir in the distal receptor zone, and the other to the duct. The duct, which is 100 μm long, has an epicuticular structure and its evacuation pore is 0.5 μm in diameter. It is surrounded by cytoplasm rich in microtubules. The epicuticular structure of the duct is resolved into fine filaments in the distal region. The ampullate glands exist only on the abdominal tergites and the pygidium. They are composed of an epicuticular receptor ampoule and a short duct connected to one secretory cell. The 2 types of glands can be placed in class 3 according to the nomenclature of Noirot and Quennedey (1974. Annu. Rev. Entomol. 19: 61–80).     

Diversity and function of male antennal glands in Cynipoidea (Hymenoptera)

The functional anatomy of antennal glands located either on the 3rd or on the 3rd and 4th antennomeres in males of several species of cynipoids was investigated. SEM observations revealed variously modified antennomeres with elevated plates, tyloids and excavated areas. In all the cases, the antennomeres are equipped with cuticular pores, corresponding internally to cuticular ducts. TEM studies showed the presence of type III integumentary glands, as classified by Noirot & Quennedey. Each glandular unit is made up of an innermost secretory cell, producing the secretion, and an outermost canal cell, producing the evacuating duct. The secretion passes through the duct and reaches the cuticular pores, concentrated in a ventro-lateral portion of the antennomere called the 'release and spread structure'. Both in Cynipidae and in Eucoilinae (Figitidae), the courtship behaviour includes a pre-copulatory phase characterized by intense antennal stroking. Bioassays in the eucoilins Leptopilina boulardi and L. heterotoma showed that these glands are the production site of a contact sex recognition pheromone, necessary for the female to accept the male.     

Glandular apparatus associated with the gastral tergal tegument in males ofStenogaster concinna andAnischnogaster laticeps (Hymenoptera: Stenogastrinae)

Summary Males ofAnischnogaster laticeps andStenogaster concinna possess clusters of glandular class III cells (Noirot and Quennedey, 1974) under the anterior edge of the 3rd gastral tergite. Their morphology is described with the help of histological sections and SEM investigation.     

Ultrastructure of the male accessory glands ofAllacma fusca(Insecta, Collembola)

The accessory glands ofAllacma fusca(L.) (Insecta, Collembola, Sminthuridae) consist of a series of secretory units that are arranged in parallel and open into the ejaculatory duct. Each unit is composed of microvillate cells stacked around a common cavity. Basal cells are involved in ion-control of fluids from the hemocoel to the cavity. The intermediate and apical cells, which have a laminar appearance and contain many microtubules, are involved in the structural integrity of the unit. Supporting cells ensheath the most apical cells. Large openings in the cuticle allow the gland secretion to flow into the ejaculatory duct lumen. These openings are protected by a porous cuticle different from that lining the epithelium of the ejaculatory duct. Conspicuous muscle fibers run along the lateroventral side of the ejaculatory duct beneath the insertion of the accessory glands. The fine structure of the accessory glands indicates that they are type I ectodermic glands as defined by Noirot & Quennedey (1974). Their function could be to control the fluidity of the material for spermatophore formation and to ensure the proper physiological conditions for spermatozoa stored in the ejaculatory duct lumen.     

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

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

Morphological characterization of female accessory sex glands of Eupolybothrus fasciatus (Newport) (Chilopoda Lithobiomorpha)

The female reproductive system of Eupolybothrus fasciatus (Newport) (Chilopoda Lithobiomorpha) includes three types of well-developed accessory glands, viz. large glands, small glands, and the periatrial gland. External morphology and the ultrastructural organization of these glands have been investigated by light and electron microscopy. The small and large glands are paired and have coiled ducts that open, respectively, into and externally to the genital atrium. By contrast, the periatrial gland is unpaired and is located on the ventral wall of the atrium into which it opens via several small canals. Ultrastructural features show that all three glands consist of two different types of cells: secretory cells and ductule cells. The secretary cells contain prominent secretory granules and are similar to a class of insect epidermal gland cells (class 3) organized as acini surrounding an extracellular lumen into which microvilli project. The granules, which have different morphological features in each gland, could be responsible for important differential functions such as producing a sexual attractant, providing a coating material that protects eggs laid on the ground, and contributing to a fluid that digests spermatophores. © 1996 Wiley-Liss, Inc.     

Fine structure of the silk gland in the mite Ornithocheyletia sp. (Prostigmata,Cheyletidae)

Silk spinning is widely-spread in trombidiform mites, yet scarse information is available on the morphology of their silk glands. Thus this study describes the fine structure of the prosomal silk glands in a small parasitic mite, Ornithocheyletia sp. (Cheyletidae). These are paired acinous glands incorporated into the podocephalic system, as typical of the order. Combined secretion of the coxal and silk glands is released at the tip of the gnathosoma. Data obtained show Ornithocheyletia silk gland belonging to the class 3 arthropod exocrine gland. Each gland is composed of seven pyramidal secretory cells and one ring-folded intercalary cell, rich in microtubules. The fine structure of the secretory cells points to intensive protein synthesis resulted in the presence of abundant uniform secretory granules. Fibrous content of the granules is always subdivided into several zones of two electron densities. The granules periodically discharge into the acinar cavity by means of exocytosis. The intercalary cell extends from the base of the excretory duct and contributes the wall of the acinar cavity encircling the apical margins of the secretory cells. The distal apical surface of the intercalary cell is covered with a thin cuticle resembling that of the corresponding cells in some acarine and myriapod glands. Axon endings form regular synaptic structures on the body of the intercalary cell implying nerve regulation of the gland activity.     

The scent glands of the male south american locust Schistocerca cancellata, an electron microscope study

The male South American locust, Schistocerca cancellata, emits a strong aromatic scent at the time of maturation. This aroma is characteristic of mature adult males living in crowded conditions. In isolated males and in females it is perceptible but faint, while nymphs emit no scent at all. Since dermal glands are numerous in the mature adult but much rarer in nymphs, young imagos and females, it seems likely that the scent is associated with these glands. The scent seems to be a maturation-pheromone, stored and released by these dermal glands. Each gland consists of a gland cell and a duct cell (type 3 gland in the classification of Noirot and Quennedey, (1974)). Each gland cell has a single end-apparatus consisting of an oblong cavity limited by the projecting tips of densely packed microvilli. A duct, dilated to form three successive bulbosities at its distal end, opens in the cavity from which it conveys the secretion to the outside. A network of fibrillar material anchors the duct to the cavity. Reproductive synchrony, beneficial in social insects, seems to be achieved in locusts by the aromatic pheromone released by the glands at the time of maturation.     

Comparing the secretory pathway in honeybee venom and hypopharyngeal glands

We provide insights into the secretory pathway of arthropod gland systems by comparing the royal jelly-producing hypopharyngeal glands and the venom-producing glands of the honeybee, Apis mellifera. These glands have different functions and different product release characteristics, but both belong to the class 3 types of insect glands, each being composed of two cells, a secretory cell and a microduct-forming cell. The hypopharyngeal secretory cells possess an extremely elongate tubular invagination that is filled with a cuticular structure, the end-apparatus, anchored against the cell membrane by a conspicuous series of actin rings. In contrast, venom glands have no actin rings, but instead have an actin-rich brush border surrounding the comparatively short and narrow end-apparatus. We relate these cytoskeletal differences to the production system and utilisation of secretions; venom is stored in a reservoir whereas royal jelly and enzymes are produced on demand. Fluorescence-based characterisation of the actin cytoskeleton combined with scanning electron microscopy of the end-apparatus allows for detailed characterisation of the point of secretion release in insect class 3 glands.     

Cutaneous eccrine glands of the foot pads of the small Madagascan tenrec (<Emphasis Type="Italic">Echinops telfairi,</Emphasis> Insectivora,Tenrecidae): skin glands in a primitive mammal

In order to find correlations between skin gland morphology and specific ethological features, the cutaneous glands of the foot pads of the primitive mammal the Madagascan tenrec, Echinops telfairi, were studied by histological and various histochemical methods as well as by electron microscopy. In the foot pads specific eccrine skin glands occurred consisting of coiled ducts and tubular secretory portions, the lumina of which were considerably wider than in primate sweat glands. The secretory tubules were composed of branched myoepithelial cells and glandular cells. The latter contained abundant mitochondria, large amounts of glycogen particles and few secretory granules as well as individual heterolysosomes and myelin bodies. The lateral cell membrane was marked by extensive interdigitations. The apical membranes of all glandular cells contained proteoglycans with sulfated and carboxylated groups containing N-acetyl-glucosamine, N-acetyl-galactosamine, galactose and mannose. The expression pattern of cytokeratins of the glandular epithelium was variable and showed similarities to that of the human eccrine glands. Tubulin, vinculin and actin were expressed in the glandular epithelium. The secretory cells showed positive reactions with antibodies against antimicrobial peptides and IgA. A positive reaction was observed with antibodies against the androgen receptor. The PCNA and TUNEL reactions indicated that the tubular skin glands of Echinops are made up of a slowly renewing tissue. We conclude that the glands fulfill several functions: production of a fluid-rich secretory product, which may prevent slipping of the foot pads on the substrate during running or climbing, secretion of antimicrobial peptides and proteins, and playing a role in thermoregulation.We thank the Fendt Foundation for financial support     

Venom gland ontogeny in Formicinae, with special reference to the pulvinate convoluted gland (Hymenoptera, Formicidae)

 The primordia of the sclerites associated with the venom gland appear in third-stage larvae. The study aims to link the structure and function of this specialised venom structure in Formicinae, together with glandular ontogeny, and puts emphasis on the relevance of the distinguished glandular subunits contributing to the final secretion. The most conspicuous changes in glandular development occur in the pharate pupa. At this stage, all subunits of the venom gland (the tubule, the convoluted gland and reservoir) are visibly present. Formation of the glandular cuticle starts around day 4 of the pupal stage. Luminal cells in the convoluted gland are provided with abundant free ribosomes and apical microvilli that remain during adult life. Stacks of granular endoplasmic reticulum are also frequently found in these cells. The convoluted gland contains relatively few scattered secretory cells, belonging to type 3 according to Noirot and Quennedey (1974), which contain electron-dense material in their extracellular spaces during adult life. These cells strongly contrast with the apparently general non-glandular nature of the convoluted gland tubule. Histochemical investigation of the secretory cells in the pulvinate convoluted gland reveals that these cells contain lipoid material, most likely to correspond with lipoids demonstrated in earlier chemical analyses. This lipoidal material in minor quantities strongly contrasts with the bulk of acid constituting the secretion. The substances produced in the convoluted gland could act as insulators, thus protecting the insect against its corrosive venom. Accepted: 28 April 1998     

Ultrastructure of male accessory glands of Chrysomya megacephala (Fabricius) (Diptera: Calliphoridae)

The ultrastructure of the male accessory glands of the blow fly, Chrysomya megacephala (Fabricius), was presented using light microscopy (LM), scanning electron microscopy (SEM), and transmission electron microscopy (TEM). A pair of accessory glands was separated at opposite sites. Morphometric results using LM yield evidenced no significant difference in the median of either length or width of the left and right glands. A significant increment in both length and width was seen to plateau between three to six days. SEM observation showed that the surface of the glands revealed a faint irregular groove pattern throughout, and it was occasionally penetrated by tracheoles. Each gland was a slender, elongated sac‐like tubule having apical rounded ends, with a slight constriction at the sub‐apical part of the gland being observed occasionally. TEM analyses of three‐day‐old males showed that the glands consisted of external capsular cells with a basement membrane underneath, glandular cells, and gland lumen. The capsular cell was flat and contained a nucleus with electron dense material in the nuclear envelope. The glandular cell, appearing as columnar, consisted of a vacuolated component that contained a large oval nucleus centrally or sub‐basally located, with dense mitochondria, numerous rough endoplasmic reticulum, and secretory vesicles containing electron‐lucent materials. In the gland lumen, the cross‐section through the middle portion revealed dense secretory materials, characterized by electron‐dense materials. Some sections revealed a large lumen where secretion accumulates within the delicate sac. The seven‐day‐old glands exhibited a remarkable change in the lumen, where the whole space contained a large amount of secretory materials, with the electron‐dense materials being characterized as similar to those observed in three‐day‐old glands. About four prominent types of secretions were observed on the basis of difference in electron‐density.     

Morphology and ultrastructure of the venom glands of the northern pacific rattlesnake Crotalus viridis oreganus

The venom gland of Crotalus viridis oreganus is composed of two discrete secretory regions: a small anterior portion, the accessory gland, and a much larger main gland. These two glands are joined by a short primary duct consisting of simple columnar secretory cells and basal horizontal cells. The main gland has at least four morphologically distinct cell types: secretory cells, the dominant cell of the gland, mitochondria-rich cells, horizontal cells, and “dark” cells. Scanning electron microscopy shows that the mitochondria-rich cells are recessed into pits of varying depth; these cells do not secrete. Horizontal cells may serve as secretory stem cells, and “dark” cells may be myoepithelial cells. The accessory gland contains at least six distinct cell types: mucosecretory cells with large mucous granules, mitochondria-rich cells with apical vesicles, mitochondria-rich cells with electron-dense secretory granules, mitochondria-rich cells with numerous cilia, horizontal cells, and “dark” cells. Mitochondria-rich cells with apical vesicles or cilia cover much of the apical surface of mucosecretory cells and these three cell types are found in the anterior distal tubules of the accessory gland. The posterior regions of the accessory gland lack mucosecretory cells and do not appear to secrete. Ciliated cells have not been noted previously in snake venom glands. Release of secretory products (venom) into the lumen of the main gland is by exocytosis of granules and by release of intact membrane-bound vesicles. Following venom extraction, main gland secretory and mitochondria-rich cells increase in height, and protein synthesis (as suggested by rough endoplasmic reticulum proliferation) increases dramatically. No new cell types or alterations in morphology were noted among glands taken from either adult or juvenile snakes, even though the venom of each is quite distinct. In general, the glands of C. v. oreganus share structural similarities with those of crotalids and viperids previously described.     

Organization of unusual idiosomal glands in a water mite, <Emphasis Type="Italic">Teutonia cometes</Emphasis> (Teutoniidae)

The unusual idiosomal glands of a water mite Teutonia cometes (Koch 1837) were examined by means of transmission and scanning electron microscopy as well as on semi-thin sections. One pair of these glands is situated ventrally in the body cavity of the idiosoma. They run posteriorly from the terminal opening (distal end) on epimeres IV and gradually dilate to their proximal blind end. The terminal opening of each gland is armed with the two fine hair-like mechanoreceptive sensilla (‘pre-anal external’ setae). The proximal part of the glands is formed of columnar secretory epithelium with a voluminous central lumen containing a large single ‘globule’ of electron-dense secretory material. The secretory gland cells contain large nuclei and intensively developed rough endoplasmic reticulum. Secretory granules of Golgi origin are scattered throughout the cell volume in small groups and are discharged from the cells into the lumen between the scarce apical microvilli. The distal part of the glands is formed of another cell type that is not secretory. These cells are composed of narrow strips of the cytoplasm leaving the large intracellular vacuoles. A short excretory cuticular duct formed by special excretory duct cells connects the glands with the external medium. At the base of the terminal opening a cuticular funnel strengthens the gland termination. At the apex of this funnel a valve prevents back-flow of the extruded secretion. These glands, as other dermal glands of water mites, are thought to play a protective role and react to external stimuli with the help of the hair-like sensilla.     

Peculiar salivary glands in a silk‐producing mite Bakericheyla chanayi (Cheyletidae)

This is the first ultrastructural investigation of salivary glands in the family Cheyletidae. In both sexes of Bakericheyla chanayi, paired acinous salivary glands and tubular coxal glands were shown to be united into the common podocephalic system. The secretory portion of the salivary gland includes medial and lateral lobes composed of the five and two cells, respectively, with clearly distinct ultrastructure. The cytoplasm of the cells is occupied by the secretory granules containing fine fibrous material. The fine structure of both cell types suggest a proteinaceous nature of their secretions. A single central process extending from the apical face of each secretory cell passes through the common acinar cavity to enter the conducting duct. A pair of intercalary cells at the base of the conducting duct links it with the secretory portion of the gland. Extending towards the acinar cavity, protrusions of intercalary cells alternate the apical regions of the secretory cells and form with them highly‐specialized contacts characterized by the apical network of microtubules and microfilaments. Two possible ways of secretion are suggested: 1) exocytosis into the acinar cavity and 2) direct passage via the central processes. The detection of axon profiles in the gland body suggests a neural control for the glandular cell function. In tritonymphs, neither secretion nor large lateral lobe cells were observed up to the pharate stage when the lateral lobe undergoes rapid differentiation. The arrangement of the acinous gland is compared to that of other arthropods. Its composition appears to be close to the class three of insect glands. The involvement of the lateral lobe cells in silk production is discussed. J. Morphol. 276:772–786, 2015. © 2015 Wiley Periodicals, Inc.     

Ultrastructural study of the mental body of Hydromantes genei (Amphibia: Plethodontidae)

The mental glands of Hydromantes genei are considered a specialized form of the urodele serous cutaneous glands. Use of a variety of techniques of maceration and digestion as well as transmission electron microscopy (TEM) and scanning electron microscopy (SEM) has shown the three-dimensional morphology of secretory and myoepithelial cells. Secretory cells are pyramidal and rest on an almost continuous layer of myoepithelial cells. The latter have a long ribbon-like body from which branch off transversal and longitudinal processes with swallow-tailed ends. Cytoplasmic processes of secretory cells, containing irregular dense vesicles, squeeze through clefts between myoepithelial cells and may reach, at some points, the basal lamina. The interstices between myoepithelium and secretory cells are extraordinarily rich in nerve endings with clear vesicles. The glandular outlets appear as elliptical stomata in the superficial layer of the epidermis and are lined by horny cells, which invaginate to circumscribe the excretory duct. The morphological results indicate that the myoepithelium of Plethodontidae mental glands differ in some respects from that of amphibian serous cutaneous glands. A double polarity for the secretory cells is also suggested. © 1993 Wiley-Liss, Inc.