Ultrastructure of the salivary glands of the planthopper,peregrinus maidis (ashmead) (Homoptera : Delphacidae)

The principal salivary gland of the planthopper, Peregrinus maidis (Ashmead) (Homoptera : Delphacidae), comprises 8 acini of only 6 ultrastructurally different acinar types. In these acini, secretory cells contain elongated vacuoles partly lined by microvilli and by microtubule bundles. These vacuoles are apparently connected with extracellular canaliculi deeply invaginated into secretory cells. Canaliculi of each acinus lead to a ductule lumen, which is lined with spiral cuticular intima, surrounded by duct cells. Striated muscle fibers, supplied with small nerve axons and tracheoles, are found in various acini of the principal gland, usually around secretory and duct cells.In the accessory salivary gland, the 2 large secretory cells contain no elongated vacuoles or canaliculi invaginations. However, in their central region, apically, these cells border a large microvilli-lined canal with its own canal cells. This canal is apparently connected with the cuticle-lined accessory duct, formed by duct cells. Nerve axons, but no muscle fibers, are found in the accessory gland and its duct. It is suggested that the system for transporting secretory material within acini of the principal gland, is basically different from that within the accessory gland.     

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.     

The labial gland in the termiteSchedorhinotermes lamanianus (Isoptera: Rhinotermitidae): Morphology and function during communal food exploitation

Summary InSchedorhinotermes lamanianus, size and cellular differentiation of the labial gland exhibit a caste-specific polymorphism. The acini of workers are composed of three distinct types of secretory cells, one of which is absent in soldiers and alates. The labial gland of workers releases a chemical signal for intraspecific communication. During the communal exploitation of a food source, labial gland secretion makes workers aggregate at gnawing sites where this signal is deposited. A newly developed bioassay is demonstrated.     

Development of the labial gland of the ponerine ant Pachycondyla obscuricornis (Hymenoptera, Formicidae) during the pupal stage

The labial gland of adult workers of the ant Pachycondyla obscuricornis is made up of many acini, each consisting of one central cell surrounded by approximately 10 parietal cells. Both cell types are associated with a system of ramified canaliculi that remove the secretion towards a ductule outside the acinus. These ductules, each associated with one acinus, fuse together and form a ramified system of ducts, ending in two paired ducts. These paired ducts widen to form a reservoir and anteriorly join into a common unpaired duct, which ends at the base of the labium. During development in the pupal stage, epithelial acini are formed first, consisting of a monolayered epithelium lining a central lumen. In these acini, one cell grows out to become the central cell, while the others will re-arrange around it to form the parietal cells. At the end of the pupal stage, the canaliculi are formed inside the acini by the central and parietal cells that secrete a lipidic substance and a cuticle. This gland type, which also occurs in some other Hymenoptera, is structurally different from the epithelial glands and the glands consisting of bicellular units, that have been traditionally distinguished until now.     

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.     

Histology,histochemistry, and ultrastructure of the harderian gland of the snake Coluber viridiflavus

Analyses of the histology, histochemistry, and ultrastructre of the Harderian gland of Coluber viridiflavus prove the gland to be compound acinar and to produce a seromucous secretion. Acinar cells (type I) contain secretory granules that are composite, consisting ultrastructurally of three distinct parts that are sharply separated. They are similar to the “special secretory granules” described in the cells of the Harderian gland of the lizard Podarcis s. sicula. Some acini of the most anterior and posterior parts of the gland are mucous. Acinar cells (type II) of this type contain secretory granules that are Alcian blue/PAS positve. At the ultrastructural level, they appear homogeneous and of low density, characteristic of mucous secretions. These mucus-secreting anterior and posterior parts of the Harderian gland may by considered as regions of intial differentiation of the anterior and posterior lacrimal galnds.     

Rosette glands in the gills of the grass shrimp,Palaemonetes pugio. I. Comparative morphology,cyclical activity,and innervation

Two types of exocrine rosette glands (called type A and type B), located in the gill axes of the grass shrimp Palaemonetes pugio, are described. The type A glands are embedded within the longitudinal median septum of the gill axes, whereas the type B glands typically project into the efferent hemolymph channels of the gill axes. Although both glands have certain common characteristics (i.e., a variable number of radially arranged secretory cells, a central intercalary cell, and a canal cell that forms the cuticular ductule leading to the branchial surface), they differ in the following respects. The type B gland is innervated, but the type A gland is not; axonal processes, containing both granular (ca. 900–1300 Å) and agranular (ca. 450–640 Å) vesicles, occur at a juncture between adjacent secretory cells and the central cell of the type B gland. The secretory cells of type A and type B glands differ in their synthetic potential and membrane specializations. These differences are more pronounced in well-developed, mature glands, most frequently encountered in larger (24–28 mm, total length) grass shrimp, than in the underdeveloped, immature glands that are most abundant in smaller (14–18 mm, total length) grass shrimp. Thus, in mature glands, the secretory cells of the type A rosette glands are characterized by extensive RER, abundant Golgi, and numerous secretory granules, whereas the secretory cells of the type B gland are characterized by extensively infolded and interdigitated basal plasmalemmas and by the presence of numerous mitochondria. In general, both types of glands exhibit increased secretory activity soon after ecdysis. The central and canal cells in both glands seem to have a role in the modification of the secreted materials. The possible functions assigned to the type A gland and the type B gland include phenol-oxidase secretion and osmoregulation, respectively.     

Morphological evidence for a probable secretory site of the male sex pheromones of Nauphoeta cinerea (blattaria,blaberidae). 2. electron microscope studies

Light and electron microscopy of the glandular epithelium of intersegmental membranes between sternites three and seven and tergites two and eight of various age groups of Nauphoeta cinerea male adults and one age group of female adults discloses differences in the epithelia of the intersternite and intertergite. The intersternal epithelium appears thicker, more glandular, and stratified. Altogether, seven cell types are recognizable, six in the male and two in the female. They are designated as types 1, 2a, 2b, 2c, 3, 4, and 5. Of these, types 1, 2a, 3, and 4 are recognizable on the sternum; types 1, 2b, and 5 on the tergum of the mature male integuments. Types 1 and 2c are found on the sternum of mature female. The cell types undergo morphological differentiation after adult emergence and show different stages of secretory activity. Type 1 are squamous cuticle-secreting cells; type 2a, 2b, and 2c are columnar-glandular and contain electron-transparent secretory vesicles of various sizes, which increase greatly in number and size in the 5-day-old adult males when the glands are most active. The vesicular size and number also differ between types 2a, 2b, and 2c cells of the same age group. The vesicles are assumed to be derived from smooth endoplasmic reticulum. The type 2 gland cells are also provided with a secretory end apparatus lined by cuticle and bordered by microvilli through which the secretion is believed to be released by exocytosis. The end apparatus leads into a cuticular ductule that opens to the surface of the cuticle as a cup-shaped receptacle, which is more conspicuous in the male intersternite. In the active gland cells, the mitochondria near the end apparatus are swollen and vacuolated. Type 3 cells are seen only on the intersternum and are believed to secrete the cuticular ductule that proceeds from the end apparatus. Type 4 cells are also recognizable only on the male intersternum and contain closely packed, electron-dense bodies, which are most numerous in mature (5-day-old) males. Type 5 cells with their dense cytoplasm are located basally in the intertergal epithelium. The functional significance of type 4 and 5 cells in the males and type 2c cells in the female is not clear. On the basis of differences in morphology, pheromone activity, and sexual behavior, it is suggested that the pheromones secreted by the intersternal and intertergal glands in the male are different, the former secreting a seducin that attracts the female to the male and the latter an “aphrodisiac” acting as a contact pheromone important in accomplishing mating.     

Ultrastructure de la glande de la spermathèque chez Phosphuga atrata L. (Coleoptera Silphidae)

Résumé Chez les Silphes et en particulier chez Phosphuga atrata, la glande de la spermathèque présente une structure particulière liée à la présence d'une intima cuticulaire tapissant la lumière de la glande. Elle comporte trois types cellulaires: les cellules sécrétrices, les cellules de l'épithélium sous-cuticulaire et les cellules-manchons. Les cellules sécrétrices de grande taille contiennent une invagination de la membrane cytoplasmique formant une «vacuole» extracellulaire bordée de microvillosités. Dans cette vacuole plonge l'extrémité, différenciée en ampoule poreuse, d'un canalicule de nature cuticulaire, qui véhicule la sécrétion jusqu'à la lumière de la glande. Le canalicule est élaboré par une cellule-manchon qui l'accompagne sur toute sa longueur sauf à son extrémité intravacuolaire.Ce type de glande, qui se retrouve chez de nombreux Insectes, y assurant des fonctions diverses (sécrétion odorifique, sécrétion de défense, sécrétion spermale, etc.), est susceptible de nombreuses variations.

---

Ultrastructure of the spermathecal accessory gland in Phosphuga atrata L. (coleoptera: silphidae)

Summary The spermathecal accessory gland in the female of Phosphuga atrata (Silphidae), exhibits a special structure which is due to the presence of a cuticular intima lining the lumen. The wall of the gland shows three cellular types: the secretory cells, the epithelial cells and the ductule carrying cells. Each large secretory cell contains a cavity formed by an invagination of the cytoplasmic membrane and lined by many microvilli. The secretory cell is connected with a cuticular ductule ending in the cavity of the glandular cell, in a porous organelle. This ductule, which carries the secretory material to the lumen, is surrounded by the ductule carrying cell.This type of integumentary gland is very common in insects, where it assumes various functions (attraction, defense, conservation of sperm, etc.) and its morphology varies considerably.

     

The terpene-producing glands of a Phasmid insect. Cell morphology and histochemistry

The defensive glands of Anisomorpha buprestoides produce the terpene toxicant anisomorphal. Each gland consists of a cuticular secretion reservoir surrounded by the secretory epithelium and the musculature which serves to compress the gland and expel the secretion. Two types of cells make up the secretory epithelium: a squamous layer next to the cuticular reservoir and a layer of larger secretory cells responsible for production of the toxicant. The microvilli-laden plasma membrane of each secretory cell is invaginated to form a central cavity. It appears that secretory products pass into the central cavity and then flow out to the gland reservoir via an efferent cuticular ductule contained within the squamous epithelial cell. Histochemical techniques demonstrate lipid reserves, carboxylic esterases, a variety of phosphatases, and an alcohol dehydrogenase, within the secretory cells. It is suggested that the lipid reserves are precursors of the terpenoid toxicant, that a mevalonic kinase has been histochemically demonstrated by the phosphatase test, and that an unusual alcohol dehydrogenase is active in the final steps of toxicant synthesis. The histochemical evidence is consistent with the hypothesis that anisomorphal is produced via the mevalonic acid pathway.     

Cellular Differentiation in a Highly Specialized Insect Secretory Organ

The colleterial glands of insects are accessory reproductive structures which produce secretions that are applied to eggs after fertilization and which serve a number of protective functions. The colleterial glands of lepidoptera are of particular interest in the study of the events of cellular differentiation because they undergo rapid development, generally during the pupal adult transformation, and contain highly specialized cells which produce large amounts of a restricted variety of secretory products. The extreme specialization of these organs facilitates parallel studies of differentiation at the biochemical and morphological level. This communication describes the changes in the ultrastructure of cells which will form the protein-secreting segment of the colleterial gland of the moth Actias luna during the period of transition from the undifferentiated state to the acquisition of secretory ability.
An initial stage of general cellular proliferation by mitosis in the presumptive colleterial tissue mass is followed by differentiation of the cells in the absence of further mitosis. Four distinctive cell types are recognized during the phase of differentiation. These types include a chitogenous cell which forms the chitin lining of the main duct, and three cells which cooperate in the formation of a secretory apparatus. Cell A forms two temporary flagella-like structures which assist in the formation of a ductule, which eventually leads from the secretory cell to the main duct. Near the end of the differentiative phase, Cell A degenerates and is phagocitized by Cell B. Cell B becomes the actual secretory element, and acquires cytoplasmic features such as extensive rough endoplasmic reticulum and Golgi apparatus which are associated with synthesis and secretion of protein. The final element, Cell C, remains associated with the ductule which it helps to construct and which traverses its cytoplasm.     

Fine structure of the intersegmental membrane glands of the sixth abdominal sternum of female Nomia melanderi (Hymenoptera,Apoidea)

The fine structure of the intersegmental glands of the sixth abdominal sternum in 1-week old females of Nomia melanderi is presented. The plasma membrane of the secretory cell is unfolded in many places and is covered by a basement membrane. The microvillous surface is invaginated to form a rather long sinuous cavity. The endoplasm is almost entirely filled by secretory granules. Many secretory granules are located close to the inner surface of the invaginated plasma membrane. The invagination contains a porous ductule, apparently of cuticulin origin, that is connected directly with the inner layer of the transport duct of the duct-forming cell. This type of arrangement allows the direct flow of the secretory substance to the outside in a continuous way. The cylindrical duct-forming cell, besides having typical cell organelles, contains a cuticular transport duct. This duct is composed of a thin cuticulin layer surrounded by a rather thick epicuticular one. The results suggest that the secretory cell has two secretory cycles. The first occurs while the gland is differentiating (at the pupal stage) and is involved in secretion of the cuticulin that forms the porous ductule. The second cycle, which starts by the beginning of nesting, is involved in the secretion of a substance that is carried to the outside via the transport duct of the duct-forming cell.     

Age-dependent changes in ultrastructure of the defensive glands of Neocapritermes taracua workers (Isoptera,Termitidae)

Protection against predators and competitors is one of the main concerns of termite colonies, which developed a specialised defensive caste, the soldiers. However, soldiers are rare or even missing in several lineages of termites, while workers often develop new defence strategies especially in soil-feeding species. Here, we describe the morphology and ultrastructure of the autothysis-associated glands of Neocapritermes taracua workers and report their age-related changes in structure. The defensive glands of N. taracua workers consist of a pair of labial and a pair of crystal glands, whose secretions mix together through autothysis. Autothysis always occurs at the line of weakness connecting the anterior parts of the crystal-bearing pouches. The crystal glands consist of groups of bicellular secretory units (secretory and corresponding canal cells) which secrete the blue crystal material into external pouches. Their secretory activity is maximal in the middle of worker life, and is considerably lower in very young and old workers. The labial glands are composed of two types of secretory cells: the central and the parietal cells. While the central cells are developed similarly to other termites and secrete proteinaceous secretion into labial gland ducts, the parietal cells develop proteinaceous granules which may eventually bud off the cells. The secretory function of parietal cells is so far unique to N. taracua and differs from other termite species in which they are only responsible of water uptake by acini. The defensive device of N. taracua is truly exceptional as it involves a new gland and a previously undescribed function for parietal cells, being a remarkable example of evolution of morphological innovation.     

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.     

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.     

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.     

Age-dependent changes in structure and function of the male labial gland in Bombus terrestris

The cephalic region of the labial gland in the buff-tailed bumblebee, Bombus terrestris, consists of numerous acini (formed by associated secretory cells and a central lumen) and connecting ducts. Age-dependent changes in secretion production (both qualitative and quantitative) are associated with changes in the amount of rough endoplasmic reticulum (RER), Golgi apparatus, and smooth endoplasmic reticulum (SER). The main secretory organelle is RER in the youngest individuals (pharate, and less-than-a-day old males), Golgi apparatus in 1-day-old males, and SER in males older than 2 days. Secretory cell death starts at 5 days of age, with maximal longevity to 10 days. Pheromone production starts immediately after eclosion, with pheromone quantities increasing until day 7. 2,3-dihydrofarnesol, the main component of the male-marking pheromone, appears in 1-day-old male glands, and reaches a maximum at 7 days of age, when its presence in the gland starts to decrease gradually. Older glands contain compounds not present in young ones. Variation in pheromone quantity and composition are reflected sensitively in the response of the queen antennae. Though queen antennae responded to gland extracts of all ages examined, maximum sensitivity was observed in response to extracts of glands 2-10 days old, while extracts of older glands gradually lose their effectiveness. Both major and minor components of the labial gland secretion extract elicited queen antennal responses suggesting that the pheromone is a multicomponent blend. Age-dependent changes in pheromone production, accumulation and tuning of pheromone activity are all synchronized approximately with male flight from the hive.     

Ultrastructure of the adenohypophysis in the larval anadromous sea lamprey,Petromyzon marinus L.

The ultrastructure of the adenohypophysis (AH) in the larval anadromous sea lamprey, Petromyzon marinus L., was examined. The AH is subdivided into three regions, the pro-, meso-, and meta-AH. Cells of the nasopharyngeal stalk extend directly beneath the pro- and meso-AH to form the ventral surface of the gland. Some cells in the pro- and meso-AH are arranged into small follicles. Each region of the AH is characterized by a single granulated (secretory) cell type. Granulated cells constitute 80–90% of the pro-AH and contain secretory granules that range from 800 to 2400 Å in diameter. Only 10–20% of the cells in the meso-AH are granulated and they contain much smaller secretory granules (400 to 1250 Å diameter) than those in the pro-AH. Granulated cells constitute 80–90% of the meta-AH and contain only a few secretory granules, ranging from 1000 to 2500 Å in diameter, and many vesicles containing either a loose flocculent or dense granular material. Nongranulated (stellate) cells are found in all regions. They are characterized by their long cell processes, abundant cytoplasmic filaments, and variable electron density. The appearance of organelles in these cells suggests they are nonsecretory. They may play a role in maintaining the structural integrity of the gland and the regulation of granule release in the pro-AH. Two types of nongranulated cells make up 80–90% of the meso-AH. Type I are stellate cells, type II may be undifferentiated cells. The functional significance of the secretory cells in the larval AH is discussed.     

Ultrastructural study of the bacillary, granular and mucoid proboscidial gland cells of Riseriellus occultus (Nemertini, Heteronemertini)

Junoy, J., Montalvo, S., Roldán, C. and García‐Corrales, P. 2000. Ultrastructural study of the bacillary, granular and mucoid proboscidial gland cells of Riseriellus occultus (Nemertini, Heteronemertini). — Acta Zoologica (Stockholm) 81 : 235–242. The ultrastructure of six types (G₅‐G₁₀) of proboscidial gland cells whose cell necks emerge independently on the epithelium surface is analysed and compared with data from other nemerteans. These types differ in cytological features, as well as in the morphology of their respective secretory granules. Secretory granules of the types G₅ and G₆ have a bacillary shape, and differ from each other based on their contents and dimensions. Secretory granules of the types G₇ and G₈ are spherical to ovoid; type G₈ gland cells are monociliated, and their secretory granules contain a paracrystalline material. Types G₉ and G₁₀ gland cells are typically goblet‐shaped; secretory granules in the type G₉ have a spherical shape, contain a homogeneous electron dense material and maintain their individuality, whereas those of the G₁₀ type are elongate and have fibrillar contents, showing a tendency to fuse before they are extruded. The mucus sheet of the proboscis is responsible for lubrication of its epithelial surface. Secretion products of type G₁₀ gland cells form the background substance of this mucus, and those of the G₅ type confer stickiness to it. Type G₉ gland cells could provide the toxic component to the mucus, and type G₇ and G₈ gland cells could be concerned with the production of enzymatic secretions.     

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.