Morphology of sternum V glands in three caddisfly species, Stenopsyche marmorata, Eubasilissa regina and Nemotaulius admorsus (Insecta: Trichoptera)

The present study describes the morphology of the sternum V gland of three caddisfly species, Stenopsyche marmorata Navas, Eubasilissa regina (McLachlan) and Nemotaulius admorsus (McLachlan), each of which belongs to a different family of the order Trichoptera, using light and scanning electron microscopy. In both sexes of these three species, the gland orifices are located on the sides of the sternum V as crescent-shaped slits, and are connected with the glandular tissue via cuticular gland ducts. The shapes of glands differ greatly among species; a slender ampullar form in S. marmorata , a flattened saccular form (horseshoe shape) in E. regina and a kidney shape in N. admorsus . The glands are composed of four essential components: large secretory cells, small reservoir cells, the lining of the reservoir and the gland duct. In S. marmorata and E. regina , additional components, muscle fibers, are present around the small reservoir cells. The secretory cells covering the whole outer surface of the gland are very large, and form many bunches in S. marmorata and E. regina , but do not form them in N. admorsus . The small reservoir cells lie inside the layer of the secretory cells and are tightly connected with the cuticular lining of the reservoir. The linings become thick cuticular ducts near the gland orifices. Histological features suggest that the secretory cells of the sternum V gland of Trichoptera belong to the type of class 3 cells in insect epidermal glands.     

Anatomy and ultrastructure of the reproductive systems ofAcarus siro (Acari: Acaridae)

Anatomy and ultrastructure of the female and male reproductive system inAcarus siro L. were investigated by light and electron microscopy. The female system consists of paired ovaries of nutrimentary type in which oogonia and oocytes are connected by bridges with a large central cell. The oviducts empty into the uterus, which passes into preoviporal duct lined bycuticle, and opening as a longitudinal slit (oviporus). An elongated accessory gland composed of one type of secretory cell is located along each oviduct. The copulatory opening occurs at the posterior margin of the body and leads, via the inseminatory canal, to the receptaculum seminis, consisting of the basal and saccular part. Both inseminatory canal and basal part of receptaculum seminis are lined by cuticle, whereas the wall of the sac is formed by cells covered only by long, numerous microvilli. The basal part of the receptaculum seminis joins the ovaries via two lumenless transitory cones.The male reproductive system contains paired testes, in which spermatogonia tightly surround the central cell. The proximal part of the paired vasa deferentia serves as a sperm reservoir, while the distal one has a glandular character. An unpaired, cuticle-lined ejaculatory duct opens into the apex of the aedeagus. The single accessory gland is located asymmetrically at the level of, or slightly posterior to, coxae IV.The structure of the genital papillae, which are topographically related to the genital opening in both sexes, is also briefly described.     

Venom and Dufour's glands of the emerald cockroach wasp Ampulex compressa (Insecta,Hymenoptera, Sphecidae): Structural and biochemical aspects

The digger wasp species Ampulex compressa produces its venom in two branched gland tubules. They terminate in a short common duct, which is bifurcated at its proximal end. One leg is linked with the venom reservoir, the other one extends to the ductus venatus. Each venom gland tubule possesses, over its entire length, a cuticle-lined central duct. Around this duct densely packed class 3 gland units each composed of a secretory cell and a canal cell are arranged. The position of their nuclei was demonstrated by DAPI staining. The brush border of the secretory cells surrounds the coiled end-apparatus. Venom is stored in a bladder like reservoir, which is surrounded by a thin reticulated layer of muscle fibres. The reservoir as a whole is lined with class 3 gland units. The tubiform Dufour's gland has a length of about 350 μm (∅ 125 μm) only and is surrounded by a network of pronounced striated muscle fibres. The glandular epithelium is mono-layered belonging to the class 1 type of insect epidermal glands. The gland cells are characterized by conspicuous lipid vesicles. Secretion of material via the gland cuticle into the gland lumen is apparent. Analysis of the polypeptide composition demonstrated that the free gland tubules and the venom reservoir contain numerous proteins ranging from 3.4 to 200 kDa. The polypeptide composition of the Dufour's gland is completely different and contains no lectin-binding glycoproteins, whereas a dominant component of the venom droplets is a glycoprotein of about 80 kDa. Comparison of the venom reservoir contents with the polypeptide pattern of venom droplets revealed that all of the major proteinaceous constituents are secreted. The secreted venom contains exclusively proteins present in the soluble contents of the venom gland. The most abundant compound class in the Dufour's gland consisted of n-alkanes followed by monomethyl-branched alkanes and alkadienes. Heptacosane was the most abundant n-alkane. Furthermore, a single volatile compound, 2-methylpentan-3-one, was identified in various concentrations in the lipid extract of the Dufour's gland.     

The exocrine glands of the male of Eldana saccharina walker (Lepidoptera,galleriinae): Intervention in precopulatory behavior

The scent apparatus of male Eldana saccharina is a glandular complex on the costal area of the forewing. It consists of two parts; glandular complex 1 is composed of five kinds of cells (epidermal cells, scale cells, glandular cells, supporting cells, duct cells); glandular complex 2 also shows five types of cells (epidermal cells, scale cells, glandular cells, duct cells, trichogen cells). The secretory products of the two parts are discharged into separate ducts which converge before opening onto the lower side of the wing. The male also has two prominent hair-pencils borne on the coremata and large secretory trichogen cells on the genital valves. Each of these exocrine gland components plays an important part in formation of the chemically complex pheromones utilized in the precopulatory behavior of the male.     

The male reproductive system and mechanism of sperm transfer in Argulus japonicus (Crustacea: Branchiura)

The histomorphology of the male reproductive system and surface morphology of the “peg-and-socket” in Argulus japonicus are described from serial sagittal and transverse sections and scanning electron micrographs. The prostate complex consists of a glandular part, a reservoir for storing the secretion, and an efferent duct opening into the ejaculatory duct. The openings of both the vas deferens and the prostate duct into the ejaculatory duct are guarded by sphincters. The ejaculatory ducts, which are lined by tall columnar epithelial cells, do not open into the cuticle-lined genital atrium but are blind-ending tubes. This observation and results obtained from observing live specimens, as well as the fact that no spermatophores are formed, suggest that semen could leave the ejaculatory duct only after puncturing of its walls. It is suggested that sperm transfer is accomplished in the following manner: during copulation contraction of the muscular walls of the vas deferens and prostate duct causes semen to be pumped into the ejaculatory duct, which is then closed off by sphincters and a high internal pressure is developed. When a spermathecal spine penetrates the walls of the ejaculatory duct, semen flows from the ejaculatory duct into the spermathecal vesicle due to the higher pressure in the ejaculatory duct. This mechanism is analogous to the sucking up of fluid with a hypodermic syringe. © 1993 Wiley-Liss, Inc.     

Evaluation of a reservoir in the main excretory duct of rat submaxillary gland.

Cannulation of salivary gland main excretory duct at its oral opening is routinely used for collecting fluid, in situ, from the luminally perfused duct, or saliva from the stimulated gland. For perfusion of the main excretory duct, in situ, or for saliva collection, rat submaxillary gland is often the organ of choice, since electrolyte transport occurs at high rates both in the whole gland and in the main excretory duct. Recently, it has been reported that there is a pouchlike dilatation of the main excretory duct at its oral end, and that this dilatation may serve as a fluid reservoir. Because of possible effects of such a reservoir on measurements of electrolyte transport by the whole gland or the main duct segment, the size and form of the reservoir have now been examined. For this, techniques of histology, radiography, and microcatherization were employed. It was found that, while the functional volume of the reservoir exceeds that of the main duct proper, the time needed for displacement of reservoir fluid by perfusate or saliva would probably be only on the order of 1-3 min at higher rates of saliva or perfusate flow. Therefore, if adequate allowance is made for equilibration time, collection of saliva or luminal perfusate by oral cannula seems justified.     

The duct system of the avian salt gland as a transporting epithelium: structure and morphometry in the duck Anas platyrhynchos

Summary The duct system of the nasal salt gland of the duck comprises central canals, secondary ducts and main ducts. The secondary and main ducts consist of a layer of columnar cells overlying a layer of small cuboidal cells. The columnar cells have complex intercellular spaces showing evidence of Na⁺ K⁺ -ATPase at the apical regions. Approximately 70% of surface area of the duct system is external to the gland. During adaptation to salt water the duct system increases in size as does the gland. Although the components of the gland of adapted ducks, including the duct system within the gland, increase in size compared with normal ducks, the percentage volume densities of the components remain similar in both categories of ducks, i.e. the duct system increases in size in proportion to the glandular tissue. The volume of the duct system external to the gland is six to seven times larger than the volume within the gland. Thus, if ductal modification of secreted fluid occurs, it will be most likely to take place in the ducts external to the gland.Total surface areas of the duct system were measured from serial sections of glands and ducts from one normal and one adapted duck. These were used to calculate possible flux rates of water and sodium across the duct epithelium, assuming the occurrence of either water reabsorption or sodium secretion. Although these flux rates are high it is shown that they are similar to calculated flux rates across the luminal surface of the secretory tubules.     

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.     

Exploring a key synapomorphy: correlations between structure and function in the sternum V glands of Trichoptera and Lepidoptera (Insecta)

The sternum V glands are a key synapomorphy that unites Trichoptera with Lepidoptera, but their functional aspects have not been analysed from an evolutionary perspective. We examine phylogenetic trends and correlations between chemical and morphological features of these glands. The most likely ancestral gland compounds are heptan‐2‐ol, 4‐hepten‐2‐one and ‐ol, nonan‐2‐one, and 6‐nonen‐2‐one and ‐ol, making pheromone production a plausible ancestral function. The most widespread gland compounds (heptan‐2‐one and ‐ol and nonan‐2‐one and ‐ol) are not known from Apataniidae + Limnephilidae (Trichoptera), which in turn uniquely produce a number of methylated 3‐ketones and their corresponding alcohols, probably functioning as pheromones. We propose a functional connection between perforated patches on sternum IV in females and a scaly/dome‐covered area around the gland openings, as well as between perforated patches and lack of Trichoptera‐type opening muscles. We also propose a functional connection between the shape of the gland reservoirs and the presence of gland reservoir musculature. The perforated patches were significantly correlated with several gland compounds that had double bonds between carbon atoms: the double bonds may lower the viscosity of the compounds, facilitating secretion through the tiny pores of the perforated patches. The production of defensive substances in Pycnopsyche (Trichoptera: Limnephilidae) is probably connected to the presence of large, compartmentalized gland reservoirs. Large glands in male Hydropsyche (Trichoptera: Hydropsychidae) are probably linked to male aggregation pheromone production. The relative sizes of sternum V gland reservoirs and fenestral gland reservoirs in female philopotamids (Trichoptera) suggest a complementary function of the two structures. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 561–579.     

Functional anatomy of scent glands in Paranemastoma quadripunctatum (Opiliones,Dyspnoi, Nemastomatidae)

The morphological organization and functional anatomy of prosomal defensive (scent) glands in Paranemastoma quadripunctatum, a representative of the dyspnoid harvestmen, was investigated by means of histological semithin sections, software‐based 3D‐reconstruction and scanning electron microscopy. Scent glands comprise large, hollow sacs on either side of the prosoma, each of these opening to the outside via one orifice (ozopore) immediately above coxa I. In contrast to the situation known from laniatorean, cyphophthalmid and some eupnoid Opiliones, ozopores are not exposed but hidden in a depression (atrium), formed by a dorsal integumental fold of the carapace and the dorsal parts of coxae I. Glandular sacs are connected to ozopores via a short duct which is equipped with a specific closing mechanism in its distal part: A layer of modified epidermal cells forms a kind of pad‐like tissue, surrounding the duct like a valve. Several muscles attached to the anterior parts of the glandular reservoir and to the epithelial pad may be associated with ozopore‐opening. The actual mechanism of secretion discharge seems to be highly unusual and may be hypothesized on the basis of corroborating data from behavioral observations, scent gland anatomy and secretion chemistry as follows: Enteric fluid is considered to be directed towards the ozopores via cuticular grooves in the surface of the coxapophyses of legs I. Then, the fluid is sucked into the anterior part of the scent gland reservoirs by the action of dorsal dilator muscles that widen the reservoir and produce a short‐term negative pressure. After dilution/solution of the naphthoquinone‐rich scent gland contents, a secretion‐loaded fluid is thought to be discharged with the help of transversal compressor muscles. This is the first detailed study on the functional anatomy of scent glands and the mechanisms of secretion discharge in the Dyspnoi. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

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.     

Queen-worker differences in spermatheca reservoir of phylogenetically basal ants

Ant queens mate when young and store sperm in their spermatheca to fertilize eggs for several years until their death. In contrast, workers in most species never mate. We have compared the histological organization of spermathecae in 25 poneromorph species exhibiting various degrees of queen-worker dimorphism. The spermathecae of both castes in all species are similar in having a reservoir connected by a sperm duct to the ovary, and a paired gland opening into this duct. The reservoir of queens typically has a columnar epithelium in the hilar region (near the opening of the sperm duct), whereas the epithelium in the distal region is cuboidal. Abundant mitochondria together with apical microvilli and basal invaginations indicate an osmoregulatory function. In contrast, the reservoir epithelium of workers is flattened throughout and lacks these transport characteristics. This single difference shows the importance of a columnar epithelium in the reservoir for sperm storage. However, our data have not revealed inter-specific variations in the development of the hilar region linked with higher fecundity. We have found no consistent differences in associated structures, such as the spermatheca gland or sperm ducts, or in the musculature between queens and workers.This work was funded by IWT, FWO, KULeuven OT and JSPS.     

Imaginal differentiation of the integument of the desert locust, Schistocerca gregaria Forsk. IV. Steps in morphogenesis of the glandular units

In the gregarious males of Schistocerca gregaria Forsk., imaginal moulting (mitosis, apolysis, new cuticle synthesis, ecdysis) is associated with the differentiation of numerous glandular units. These units, involved in a sexual, excitatory pheromone secretion are at that time composed only of a basal glandular cell and an apical duct cell. Each glandular unit originates from an isogenic group of cells of which the four elements (tetrade) are disposed on two levels. At each level a principal cell and an accessory one may be recognized. The lower accessory, or ciliary, cell shows, at the time of apolysis, both a strong cytoplasmic protrusion and a typical ciliary formation. This formation associated with a diplosome goes through the duct cell and ends up in the exuvial space. It makes a inner mould; arranged around it are epicuticular materials characteristic of the duct wall; then it disappears. The strong cytoplasmic protrusion also retracts thus allowing a glandular reservoir to form. A glandular cell may be recognized at an early stage owing to its R.E.R. development. The upper accessory cell strengthens the duct cell and secretes junctional cuticle between the duct and general cuticle. Accessory cells, after the imaginal moult do not degenerate but acquire epithelial cell characteristics. The duct has a dual origin : the receptive part is secreted by the ciliary cell and the vector part by the duct cell. The organization and stages of morphogenesis of the glandular unit are discussed and compared to those of other apterygote or pterygote insects.     

The sting bulb gland in myrmecia and Nothomyrmecia (Hymenoptera : Formicidae): A new exocrine gland in ants

A new exocrine gland has been discovered within the sting of the endemic Australian ants of the genera Myrmecia and Nothomyrmecia (Hymenoptera : Formicidae). It consists of approximately 20 secretory cells with their accompanying duct cells, located between the ducts of the venom and Dufour glands in the proximal part of ths sting bulb, hence my suggestion to designate it as the sting bulb gland. Ultrastructural examination reveals the development of both granular and smooth endoplasmic reticulum in the glandular cells, which possibly may indicate the elaboration of a rather complex secretion. Although the function of the gland remains unknown, its exclusive presence in these ants provides another argument for a closer phylogenetic relationship between both genera than is reflected by their actual classification.     

The spermathecal complex in Ips typographus (L.): Differentiation of the spermathecal gland related to age and reproductive state

The spermathecal complex of the bark beetle, Ips typographus, comprises the following elements: spermathecal duct, spermatheca and spermathecal gland. The spermathecal duct connects the vagina and the spermatheca and consists of a cuticular tube surrounded by an epithelial layer and circular muscles. The spermatheca is bottle-shaped and has a cuticle-lined lumen. Muscles are attached to both ends of the spermatheca. The spermathecal gland which is connected to the spermatheca possesses three cell types: glandular, hypodermal, and ductule. The glandular cells have different structural characteristics depending on the age and reproductive state of the females. After the emergence of the brood, two different kinds of secretory material are present in the glandular cells. There is evidence that one type of secretion is emitted during the first few days after brood emergence, while the other type accumulates to be secreted during later stages.     

Genital system development of Williamia radiata (Gastropoda, Siphonariidae)

The adult anatomy and development of the genital system of the limpet Williamia radiata (Siphonariidae, Gastropoda) was investigated by means of 3D computer reconstruction and visualization of serial resin and paraplast light microscopical sections. As is characteristic for siphonariid species, the adult genital system consists of a single duct, the spermoviduct leading from the nidamental glandular system to the anteriorly located genital atrium with opening. The copulatory complex as well as the bursa copulatrix also open into the common genital atrium. The genital system develops from three separate anlagen. The posterior one appears first at a body length of 0.7 mm and gives rise to the ovotestis and part of the hermaphrodite duct. The nidamental glandular complex, the fertilization pouch-spermatheca complex, part of the hermaphrodite duct, the posterior part of the spermoviduct, and the bursa copulatrix develop later from the pallial anlage. Finally, the anterior anlage is formed on the right side of the head and gives rise to the genital atrium, the copulatory complex, and the anterior spermoviduct. This formation of the genital system from three, locally separated anlagen, differs strikingly from that of most other species of the Euthyneura. In both the Nudibranchia (Opisthobranchia) and the Stylommatophora (Pulmonata) development proceeds from a single site. We regarded this as a secondary condition as a result of derived features like heterochronies in development in these taxa. Comparison of development with that of other species of the Pulmonata allows conclusions on homology. The homology of the bursa copulatrix within the pulmonates is confirmed. The two separate chambers inside the spermoviduct of W. radiata correspond to oviduct and vas deferens of the freshwater Basommatophora.     

Ducts of the labral glands of Leptestheria dahalacensis (Crustacea: Branchiopoda: Spinicaudata)

Inside the labrum of Leptestheria dahalacensis are situated three types of large epidermal gland cells, whose ducts open onto the outer dorsal surface of the labrum. SEM revealed that the thin ducts of the A-type gland cells open out behind the epipharynx at the end of small, conically shaped protuberances, the two paired ducts of the B-type gland cells lead into the distal portion of the labrum, and the external opening of the single duct of the C-type gland cells lies on the dorsal lobe of the labrum. The ducts of the three different gland cell types have the same fundamental constitution, but vary in diameter. Each secretory unit consists of a pair of gland cells (A, B, or C) and a secretory duct. The duct is formed by ring-shaped folding of one anteroposteriorly elongated epidermal cell (duct cell), whose ends adhere closely to one another. A further ring-folded epidermal cell (accessory cell), but flattened in shape, is interposed, like a sleeve-connection, between the gland cells and the duct cell. The reservoirs of gland cells open into the lumen of the duct. Discontinuous deposits of highly electron-dense matter are present on the plasma membrane of the accessory cell delimiting the initial part of the duct lumen, while the plasma membrane of the duct cell facing the lumen is cuticularized. The cytoplasm of the accessory cell, on examination by TEM, appears quite similar to that of the duct cell, except for the different distribution and greater abundance of microtubules. Similarly organized tricellular tegumental glands also commonly occur in other Crustacea, both Malacostraca and non-Malacostraca. Possible functions of secretions from the three different types of gland cells present in the labrum of L. dahalacensis are discussed.