Ultrastructural modifications of the glandular epithelium of the receptaculum seminis in Thermobia domestica (Insecta: Thysanura) during the moulting period

Summary The wall of the receptaculum seminis of Thermobia domestica is composed of numerous glandular units, each with four enveloping cells (denoted 1 to 4) separated by ordinary epithelial cells and associated with a cuticular apparatus. During the moulting periods, which continue to occur in the adult stage, these cells undergo a series of transformations. Just before apolysis there is a dedifferentiation of numerous cytoplasmic organelles, but no mitosis has been observed. When the intima lifts off, the apical system of each glandular unit, i.e. the distal parts of the C2 and C3 cells surrounding the end apparatus, is also eliminated. Then at the apex of each glandular unit, a new ductule is formed in the cavity of which a long ciliary process grows up from cell C1. Finally comes the phase of cuticle formation, i.e., epicuticle for the ductules, epi-and endocuticle for the intima lining the central cavity of the receptaculum. Various cell types participate in secretion of cuticle, the ciliary cells (C1) being responsible for the formation of the porous end apparatus. At ecdysis almost all of the new intima has been secreted and the apical systems are once more differentiated. These transformations are compared with those recently described in other exocrine glands of arthropods, e.g., tegumentary glands and accessory glands of the genital ducts.     

A light and electron microscopical study of the spermathecae and ventral receptacle of Anastrepha suspensa (Diptera: Tephritidae) and implications in female influence of sperm storage

Female insects with multiple sperm storage organs may potentially influence patterns of paternity by differential storage of sperm from competing males. The Caribbean Fruit Fly, Anastrepha suspensa, stores sperm differentially with respect to its three spermathecae. To understand the mechanisms and processes responsible for patterns of sperm storage and use in A. suspensa, details of the fine structure of female sperm storage organs were resolved by UV-light microscopy, confocal microscopy, tissue sectioning, and scanning and transmission electron microscopy. Structures not previously described for this species include a ventral receptacle for sperm storage and osmoregulation, a conical-shaped valve at the junction between the spermathecal capsules and their ducts, laminar and granular secretions, secretions from the signum, hemocytes surrounding the spermathecae, and spermathecae with sclerotized, hollow projections that terminate in single glandular cells. The independent organization of sperm storage organs, spermathecal ducts, associated musculature, gland cells, and innervation offer possible mechanisms by which sperm movement may be influenced by females. The implications of these structures for insemination and fertilization events are discussed.     

黄胫小车蝗受精囊的亚显微结构

利用组织学方法,观察了黄胫小车蝗Oedaleus infernalis 受精囊的显微与亚显微结构。结果表明,黄胫小车蝗受精囊为单个,由高度卷曲的受精囊管和蚕豆状的端囊构成。受精囊壁主要由表皮层、上皮层、基膜和肌肉层构成;上皮层包含上皮细胞、导管细胞和腺细胞。上皮细胞在靠表皮层的边缘有大量的微绒毛,两相邻上皮细胞的细胞膜相互嵌入,并有细微的突起延伸在导管细胞及腺细胞之间,直到基膜,达基膜处的上皮细胞膜折叠,与腺细胞膜的折叠,一起形成迷宫样的指状突起,附着在基膜上。导管细胞有一个较大的核和分泌导管,连接于腺细胞的细胞腔和受精囊腔,将腺细胞中分泌物运输到受精囊腔中。腺细胞具有典型的分泌细胞特征： 含发达内质网、高尔基复合体及不同大小的囊泡。肌肉层位于受精囊最外层,附在基膜上。在受精囊不同部位的结构有差异。在交配前和交配后,受精囊腺细胞的亚显微结构也有差异。     

Ultrastructure of the phallic glands of the firebrat,Thermobia domestica (packard) (Thysanura : Lepismatidae)

The exocrine glands located in the penis of Thermobia domestica (Thysanura : Lepismatidae) are composed of about 100 distinct units, each containing several cell types: one large secretory cell with an apical reservoir; 2 groups of envelope cells, an inner group of 2 superimposed cells, and an outer group of 4 cells arranged in a ring, and also 2 basal cells, called ciliary cells owing to their elongated processes, which look like the dendrite of a sensory cell. Each functional unit includes cuticular differentiations: a tubular bristle, fixed on a small tubercle; and a long “internal” ductule communicating basally with the reservoir of the glandular cell and opening distally at the tip of the bristle. A study of the modifications affecting the phallic glands during moulting, shows that the inner envelope cells deposit the cuticle that forms the ductule, the outer envelope cells elaborate the cuticule of the tubercle, while a temporary distal projection of only one of these cells ensures the formation of the bristle. In addition, a lengthening of the outer dendritic segment of the 2 ciliary cells takes place before ductule formation, but this segment partially degenerates after ecdysis. These findings are compared with data already obtained on the morphogenesis of other insect integumental glands. In T. domestica, the secretion of the phallic glands is presumed to be used, during the mating sequences, for spinning fine threads before spermatophore deposition.     

An electron microscopic study of the spermathecal complex of virgin Aedes aegypti mosquitoes

Virgin mosquitoes were studied with the electron microscope. Spermathecal duct walls contain cuticle, epithelium, and a richly innervated spiral muscle; myocytes are linked by desmosome-like attachment plaques to the underlying epithelium. Periductal cells along upper portions of the ducts have a large secretory droplet within a highly irregular extracellular lacuna and are attached to a long secretory ductule through which finely granular material is delivered to the duct lumen and this enters the spermathecae. Basal gland cells of spermathecae have short ductules containing secretion in virgins. Secretory material in spermathecae of virgins does not form a complete internal membrane.     

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.     

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.     

Female genital system of the folding-trapdoor spider Antrodiaetus unicolor (Hentz, 1842) (Antrodiaetidae, Araneae): ultrastructural study of form and function with notes on reproductive biology of spiders

The genitalia of the female folding-trapdoor spider Antrodiaetus unicolor are characterized by two pairs of spermathecae that are arranged in a single row and connected to the roof of the bursa copulatrix. Each single spermatheca is divided into three main parts: stalk, bowl, and bulb, which are surrounded by the spermathecal gland. The epithelium of the spermathecal gland is underlain by a muscle meshwork and consists of different types of cells partly belonging to glandular cell units (Class 3 gland cells) that extend into pores in the cuticle of the stalk and bowl. Interestingly, the bulb lacks glandular pores and is characterized by a weakly sclerotized cuticle. This peculiarly structured bulb probably plays an important role in the discharge of the sperm mass. It is suggested that by contraction of the muscle layer the sperm mass may be squeezed out, when the bulb invaginates and expands into the spermathecal lumen, pushing the sperm to the uterus lumen. Each glandular unit consists of usually one or two central secretory cells that are for the most part surrounded by a connecting cell that again is surrounded by a canal cell. The canal cell, finally, is separated from the other epithelial cells (intercalary cells) located between the glandular units by several thin sheath cells that form the outer enveloping layer of the unit. The secretions are released through a cuticular duct that originates proximally between the apical part of the connecting cell and the apical microvilli of the secretory cells and runs into a pore of the spermathecal cuticle. The glandular products of the Class 3 gland cells likely contribute to the conditions allowing long-term storage of the spermatozoa in this species. Details regarding the ovary, the uterus internus, and the uterus externus are reported. Most of the secretion that composes the chorion of the egg is produced in the ovary. Glandular cell units observed in the uterus externus differ structurally from those in the spermathecae and likely play a different role. Finally, we briefly discuss our results on the female genitalia of A. unicolor in the light of knowledge about the reproductive biology of spiders.     

Cytodifferentiation in the accessory glands of Tenebrio molitor. III. Fine structure of the spermathecal accessory gland in the pupa

To establish indices for studying the hormonal control of differentiation of the accessory reproductive glands of insects, the ultrastructural development of the spermathecal accessory gland (SAG) of female mealworm beetles has been analyzed. Over the 9 days between adult and pupal ecdysis, the SAG transforms from a stubby sac of columnar epithelium into an elongate cylindrical gland, lined with cuticle, and containing several distinct types of differentiated cells. The first phase of pupal differentiation is one of cell division and overall gland morphogenesis which lasts 3--4 days; at its close, two populations of cells can be distinguished. One of these populations will produce the cuticular ductules while the other will yield the three-cell secretory units or organules. In the second phase which lasts 2 days, the three cells of each organule become wrapped around one another and then the innermost puts out a pseudocilium and retracts within the next ensheathing cell. In the third phase which lasts 4 days, the cuticles of the axial duct, of the efferent ductule, of the vestibule upon which the ductules converge, and of the end apparatus, are deposited. The ciliary process degenerates, and after ecdysis, the secretory cells undergo peak differentiation.     

Structure and function of the spermathecal complex in the phlebotomine sandflyPhlebotomus papatasi Scopoli (Diptera: Psychodidae): I. Ultrastructure and histology

Females of phlebotomine sandflies (Diptera: Psychodidae) possess highly variable spermathecae that present several important taxonomic characters. The cause of this diversity remains a neglected field of sandfly biology, but may possibly be due to female post-mating sexual selection. To understand this diversity, a detailed study of the structure and function of the spermathecal complex in at least one of the species was a prerequisite. Using scanning and transmission electron microscopy, described here is ultrastructure of the spermathecal complex in the sand fly,Phlebotomus papatasi Scopoli. The spermathecal complexes are paired; each consists of a long spermathecal duct, a cylindrical spermathecal body, and a spherical spermathecal gland. Muscle fibres, nerves, tracheoles, and vascular sinuses connect the spermathecal body and duct through the epithelial layers. Spermathecal gland is formed by a typical insect epidermis and consisting of an epithelial layer of class-1 epidermal cells and elaborate glandular cells of class-3 epidermal cells, each having both receiving and conducting ductules (i.e. “end apparatus”) and a “cytological apodeme”, which is a newly described cell structure. The spermathecal body and duct are lined by class-1 epidermal cells and a cuticle, and are enveloped by a super-contracting visceral muscular system. The cuticle consists of rubber-like resilin, and its fibrillar arrangement and chemical nature are described. A well-developed neuromuscular junction exists between the spermathecal gland and the spermathecal body, which are connected to each other by a nerve and a muscle. The spermathecal complexes of the sandfly are compared with those of other insect species. The physiological role and possible evolutionary significance of the different parts of spermathecal complex in the sandfly are inferred from the morphology and behaviour. Post-mating sexual selection may be responsible for the structural uniqueness of the spermathecal complex in phlebotomine sandflies.     

Modifications ultrastructurales des glandes vésiculaires des Machilidae (Insecta,Thysanura) au cours des cycles de mue

Summary During the period between apolysis and ecdysis, the vesicular glands show many important transformations which affect not only the cuticular ductules, but all the cells. The cytoplasm of the glandular cells undergoes a partial autolysis, whereas other parts of the cells present a high secretory activity. Immediately after the apolysis the cellular reservoir empties and disappears almost completely; soon after, refills with secretion. The most interesting transformations concern each ciliary cell, always associated with a glandular cell. In the first phase of the moulting cycle, the dendrite of the ciliary cell grows a ciliumlike extension (= distal region of the dendrite), which penetrates into the corresponding ductule; the new intima of this ductule is laid around the cilium. At the same time, the proximal region of the dendrite forms a circular fold around the base of the cilium and begins to secrete a material which will form the end apparatus. This latter is finished during the second phase of the cycle. The third phase is characterized by the degeneration of the distal region of the dendrite and the circular fold. Thus, the end apparatus is not a secretion of the ductule-carrying cell, but of the ciliary cell. At the end of the moulting period, just before ecdysis, the vesicular gland again takes the structure characteristic of the intermoult: the reservoir of the glandular cell is very large; the cuticular apparatus is almost formed; the dendrite of the ciliary cells shows, at its apex, a short cilium (= ciliary region s. str. + short distal region) surrounded by microvilli, free in the secretion of the reservoir.     

Rosette glands in the gills of the grass shrimp Palaemonetes pugio. II. Premolt ductule reformation: Replacement of ciliary processes by cytoplasmic processes in relation to gland maturation

The events associated with premolt reformation of the cuticularized ductule in the underdeveloped (immature) branchial rosette glands, which are common in the gills of small (14–18 mm, total length) grass shrimp, are described and contrasted with the events of ductule reformation in the fully developed (mature) resette glands most common in larger shrimp. In immature rosette glands, two ciliary processes emerge from each of the component secretory cells and ascend into the basal luminal region of the old ductule. Subsequently a new ductule is formed around the old ductule, and the ciliary processes disappear, either because of degeneration or retraction. The transitory ciliary processes appear to prevent the old ductule from collapsing during the formation of a new ductule. Such transitory ciliary processes, however, are not found in association with premolt ductule reformation in the mature rosette glands; in their place are seen a number of microvilli-like cytoplasmic processes, which emanate from the apices of the secretory cells and from the channels of the central cell. These cytoplasmic processes in mature glands, like the ciliary processes in immature glands, are transitory and appear to prevent the collapse of the old ductule. Cytoplasmic processes comparable to those in mature glands, but relatively few in number and originating only from the secretory cells, are seen together with ciliary processes in some immature glands. The relative abundance of cytoplasmic processes in the mature glands, coupled with the observation that transitory ciliary processes occur in immature glands but not in mature glands, suggests that, during glandular maturation, transitory ciliary processes are replaced by transitory cytoplasmic processes.     

Internal reproductive organs of the female humpbacked fly,Megaselia scalaris Loew (Diptera : Phoridae)

The female reproductive system of the humpbacked fly Megaselia scalaris Loew (Diptera : Phoridae) was examined in whole mount preparations and serial sections. The system includes 2 ovaries, paired lateral oviducts, a common oviduct, and a genital chamber, opening externally through a gonopore, anteriad and ventrad to the anus. The ducts of the 2 accessory glands open independently into the dorsal region of the genital chamber. The terminal duct of a 2-armed spermatheca joins the right posterior and ventral wall of the genital chamber, immediately inside the gonopore. Passing dorally, the spermathecal duct lies immediately ventral to the duct of the right accessory gland. A short distance posteriad, it divides into two branches, each supplying an arm of the spermatheca. The genital chamber extends both anteriorly and posteriorly from its junction with the common oviduct, creating anterior and posterior compartments. In the right lateral wall of the genital chamber, a distinctive loop-shaped thickening (plate) resembles a darkened thread when it is observed through the integument. Features likely to have taxonomic utility include the posterior and ventral location of the terminal portion of the spermathecal duct; and the asymmetrically arranged, loop-shaped plate.     

Ultrastructure of an exocrine dermal gland in the gills of the grass shrimp,Palaemonetes pugio: Occurrence of transitory ciliary axonemes associated with the sloughing and reformation of the ductule

Exocrine dermal glands, comparable to the class 3 glandular units of insects, are found in the gills of the grass shrimp, Palaemonetes pugio. The dermal glands are composed of three cells: secretory cell, hillock cell and canal cell. Originating as a complex invagination of the apical cytoplasm of the granular secretory cell, a duct ascends through the hillock and canal cells to the cuticular surface. The duct is divisible into four regions: the secretory apparatus in the granular secretory cell, the locular complex, the hillock region within the hillock cell and the canal within the canal cell. A tubular ductule is contained within the latter two regions. As the ductule ascends to the cuticular surface, its constitution gradually changes from one of a fibrous material to one which possesses layers of epicuticle. During the proecdysial period, the ductule is extruded into the ecdysial space and this is followed by the secretion of a new ductule. Temporary ciliary structures, located near the secretory apparatus of the secretory cell, are associated with the extrusion and reformation of the ductule. Characterized only by a basal body and rootlets throughout most of the intermolt cycle, the ciliary organelles give rise to temporary axonemic processes which ascend through the ductule toward the ecdysial space at the onset of proecdysis. Subsequently, the old ductule is sloughed off and a new ductule is reformed around the ciliary axonemes. Following this reformation, the ciliary axonemes degenerate. The function of cytoplasmic processes, derived from the apical cytoplasm of the secretory cell, is also discussed.     

Genital structures in the entelegyne widow spider Latrodectus revivensis (Arachnida; Araneae; Theridiidae) indicate a low ability for cryptic female choice by sperm manipulation

The female genital structures of the entelegyne spider Latrodectus revivensis are described using semithin sections and scanning electron microscopy. Apart from the tactile hairs overhanging the opening of the atrium, the contact zones of the female epigynum are devoid of any sensilla, indicating that the female does not discriminate in favor or against males due to their genital size or stimulation through copulatory courtship. The dumb-bell shape and the spatial separation of the entrance and the exit of the paired spermathecae suggest that they are functionally of the conduit type. Not described for other entelegyne spiders so far, the small fertilization ducts originating from the spermathecae of each side lead to a common fertilization duct that connects the spermathecae to the uterus externus. During oviposition, it is most likely that spermatozoa are indiscriminately sucked out of the spermathecal lumina by the low pressure produced by the contraction of the muscle extending from the epigynal plate to the common fertilization duct. As no greater amounts of secretion are produced by the female during oviposition, and no activated sperm are present within the female genital tract, the secretion produced by the spermathecal epithelium does not serve in displacement or (selective) activation of spermatozoa. These findings suggest that female L. revivensis are not able to exert cryptic female choice by selectively choosing spermatozoa of certain males.     

Structural organization of the copulation site in the medfly Ceratitis capitata (Diptera: Tephritidae) and observations on sperm transfer and storage

The copulation site of the medfly Ceratitis capitata was investigated at anatomical and ultrastructural levels. It consists of the anterior vagina, with a ventral fertilization chamber and a dorsal insemination pocket into which the two spermathecal ducts open. The fertilization chamber is an organ comprised of a number of alveoli that in virgin females are filled with a filamentous secretion, whereas in mated females contain sperm bundles. Through study of the internal morphology of the aedeagus, its position in the anterior vagina, and the direct observation of sperm transfer and storage, we confirmed that sperm are ejaculated through two gonopores at the top of the distiphallus and another at the base of the genital rod. The sperm flow dorsally into the insemination pocket and ventrally into the fertilization chamber. During copulation, the two spermathecae and the fertilization chamber are progressively filled with spermatozoa.     

Evidence of a neural loop involved in controlling spermathecal contractions in Locusta migratoria

The control of spermathecal contractions in Locusta migratoria via a neural loop was demonstrated using mechanical stimulation and electrophysiological recordings. Extracellular electrophysiological recordings from the receptaculum seminis nerve (N2B2), which innervates the spermathecal sac, were conducted during mechanical stimulation of the genital chamber sensory cells. Activation of the genital chamber sensory cells, using a glass probe approximating the shape and size of an egg, was found to increase the action potential frequency and initiate bursts of action potentials if a tonic frequency of action potentials was present prior to stimulation. If the motor pattern initially consisted of bursts of action potentials, then mechanical stimulation of the genital chamber sensory cells resulted in an increase in firing frequency, in most preparations, with the bursting remaining. Removal of the probe from the genital chamber always returned the motor activity to that noted prior to sensory cell stimulation. Simultaneous electrophysiological recordings from both the left and right receptaculum seminis nerves (N2B2) revealed that the bursts of action potentials were coordinated, although individual action potentials were not coupled one to one. Activation of the genital chamber sensory cells also resulted in increases in spermathecal contraction frequency, an effect which was coordinated with the changes in motor activity. It is proposed that an egg in the genital chamber activates the sensory cells resulting in increases in spermathecal contraction frequency and the subsequent release of spermatozoa onto the micropyle of the egg for fertilization.     

The morphogenesis of spermathecae and spermathecal glands in Drosophila melanogaster

Sperm storage in female insects is important for reproductive success and sperm competition. In Drosophila melanogaster females, sperm viability during storage is dependent upon secretions produced by spermathecae and parovaria. Class III dermal glands are present in both structures. Spermathecal glands are initially comprised of a three-cell unit that is refined to a single secretory cell in the adult. It encapsulates an end-apparatus joining to a cuticular duct passing secretions to the spermathecal lumen. We have examined spermatheca morphogenesis using DIC and fluorescence microscopy. In agreement with a recent study, cell division ceases by 36 h after puparium formation (APF). Immunostaining of the plasma membrane at this stage demonstrates that gland cells wrap around the developing end-apparatus and each other. By 48–60 h APF, the secretory cell exhibits characteristic adult morphology of an enlarged nucleus and extracellular reservoir. A novel finding is the presence of an extracellular reservoir in the basal support cell that is continuous with the secretory cell reservoir. Some indication of early spermathecal gland formation is evident in the division of enlarged cells lying adjacent to the spermathecal lumen at 18 h APF and in cellular processes that bind clusters of cells between 24 and 30 h APF.