Functional anatomy of male copulatory organs of Pomacea canaliculata (Caenogastropoda, Ampullariidae)

This study was aimed to investigate the functional morphology of copulation and sperm transfer in the invasive snail Pomacea canaliculata. Three-dimensional renderings of the male copulatory apparatus were made and showed elaborate systems for innervation and for hemolymph supply and drainage. A key component of the male copulatory apparatus is the penial sheath, which shows three specialized glands; the medial and distal glands may participate in adherence to the mantle cavity wall of the female during copulation. The outer gland has an epithelium composed of columnar cells with branched microvilli, mucous goblet cells and large granular secretory cells containing intragranular crystalloids, which produce an exocrine secretion during copulation. The interaction of male/female copulatory organs was studied in dissections of snap-frozen pairs. Sperm are left in the sperm pit, at the end of the pallial spermiduct. Afterwards, the muscular action of the penial bulb takes the sperm up to the vermiform penis, which slides from the penial pouch into the central groove of the penial sheath, and it later emerges through a T-shaped sulcus of this structure and enters the female vagina. Then, it climbs through the capsule duct, and its tip reaches the proximity of the seminal receptacle. A model of copulation and sperm transfer is presented on the basis of the new findings and on published literature.     

Ultrastructure of the male reproductive system and spermatophore formation in Labidocera aestiva (Crust.: Copepoda)

Summary The male reproductive system of Labidocera aestiva produces a flask-shaped spermatophore connected to a chitin-like coupling apparatus. As immature spermatozoa leave the anterior region of the testis, they pass through the lumen of a long, sinuous duct composed of a ductus deferens and seminal vesicle. Ultrastructural examination of the ductus deferens reveals a highly glandular, columnar epithelium. The cells contain arrays of rough endoplasmic reticulum and abundant, well-developed Golgi complexes. This region produces and releases into the lumen, a flocculent substance and two granular secretions that constitute the seminal fluid. In its terminal part, the ductus deferens synthesizes another secretion that forms the spermatophore wall enclosing the spermatozoa and seminal fluid. Final synthesis of the spermatophore wall occurs within the thin-walled seminal vesicle, although this region functions primarily as a storage organ. Contiguous to the seminal vesicle is an elongate, highly glandular spermatophore sac. The chitin-like coupling apparatus, which functions to attach the spermatophore to the female, is formed in the anterior region of the sac by secretions from eight cell types. The posterior region of the sac stores the flask-shaped spermatophore and produces secretions that aid ejaculation of the entire spermatophore complex.Contribution No. 236, Harbor Branch Foundation, Inc.     

New insights into the functional morphology of the male copulatory apparatus of bullid gastropods

In this study, semi-thin sections stained with histochemical techniques and transmission electron microscopy were used to obtain new data about the morphology and function of the male copulatory apparatus of the cephalaspidean gastropod Bulla striata. The apparatus comprises a vestibule, a penial papilla and a prostate consisting of a coiled unbranched tube ending in a blind caecum. The penial papilla and the coiled tubular prostate are enclosed by a muscular sheath, which is continuous with the muscular tissue of the vestibule. The epithelium lining the lumen of the vestibule is formed by ciliated and mucus-secreting cells. Two new types of subepithelial secretory cells were discovered in this region. The penial papilla is a muscular structure without secretory cells in the epithelium lining the narrow lumen. The tubule that constitutes the prostate possesses a muscular wall and can be divided in three distinct regions: a non-secretory duct connected to the penial papilla, a glandular region rich in large secretory cells and the terminal caecum containing just a few small secretory cells. In the terminal blind caecum, the muscular sheath is fused with the muscular wall of the tubular prostate. Large numbers of spermatozoa were found in the glandular region and in the terminal caecum of the prostate. A new functional mechanism is proposed to explain penial eversion during copulation. This differs from a previous hypothesis in two main aspects: (1) existence of a permanent penial papilla in mature animals acting as a functional penis and (2) functional role of vestibule during copulation, which everts and surrounds the penial papilla, while the latter protrudes outwards.     

Functional morphology of the mammiliform penial glands ofLittorina saxatilis (Gastropoda)

Summary The functional morphology of the mammiliform penial glands ofLittorina saxatilis has been investigated with both light and electron microscopy. These penial glands line the ventral edge of the penis and orient with the female mantle during copulation. Secretions are released from the penial glands to this interface where they probably function in adhesion. The penial gland secretions comprise heterogeneous granules as well as apocrine and mucous secretions. The heterogeneous granules are produced in separate multicellular glands arranged in a series of lobes that lie outside a thick smooth muscle layer enclosing the lumen. Each glandular lobe is surrounded by a thin layer of smooth muscle. Secretions are transported in individual cellular processes that pass through the thick smooth muscle layer and empty into the lumen. Surrounding the lumen is an epithelium containing apocrine secretory cells as well as occasional goblet-type, mucous cells. The combined action of the muscles forces secretions out of the lumen through the penial papilla, onto the external surface of the mammiliform penial gland. Longitudinal muscles extend into the penial papilla enabling its protrusion or retraction. Retraction of the penial papilla following secretion release is thought to create negative pressure beneath the penial gland producing suction adhesion. The visco-elastic properties of the penial gland secretion are qualitatively different from foot mucus and may represent specialization to an adhesive function.     

Histological,histochemical, and ultrastructural investigation of the male copulatory apparatus of Haminoea navicula (Gastropoda,Cephalaspidea)

Due to its biological and systematic importance, the morphology and function of the male copulatory apparatus of Haminoea navicula, a Cephalaspidea gastropod mollusk, was investigated by light and electron microscopy. These systems are poorly understood in haminoids, but are often used in the taxonomy of the genus. In H. navicula, the male copulatory apparatus comprises the penis within a penial sheath, a seminal duct and the prostate with two lobes. The penis is a muscular structure with a tip covered by spikes formed by muscular cells. The penial sheath consists of muscular tissue folds lined by an epithelium. Below this epithelium, polysaccharide‐secreting cells and pigment cells were observed. A large number of vacuolar cells were found below the ciliated epithelium of the seminal duct. The proximal lobe of the prostate was formed by tubules that could be divided in basal, middle and apical zones, containing cells that secrete polysaccharides and proteins. The tubules of the prostate distal lobe contained a single type of secretory cells with vesicles that were stained by histochemical techniques for detection of polysaccharides and proteins. Ciliated cells were present along the tubules in both lobes of the prostate. This study revealed a complex prostate with five types of secretory cells, which secrete substances that should be involved in the maintenance and eventually also in the maturation of spermatozoa. A comparison with previous publications, shows that the male copulatory apparatus can differ substantially among cephalaspideans, even between H. navicula and non‐European species attributed to this genus.     

Morphological effects of tributyltin (TBT) in vitro on the genital system of the mesogastropodaLittorina littorea (Prosobranchia)

This ultrastructural study investigates the pathological changes in the penial, the sperm groove and the glandular cushion epithelium in maleLittorina littorea (Mesogastropoda) related to TBT (tributyltin) contamination. The results are compared with those onOcinebrina aciculata (Neogastropoda), which shows a wide range of cell changes in the penis epithelia of male and imposex affected females. The investigation of the different penis epithelia ofL. littorea revealed that the cells analysed show a low sensibility towards TBT. The cells display normal metabolism. Certain atypical structures like swelling microvilli and cristae which tend to lie parallel to the long axis in the mitochondria, were detected as pathological effects.     

The fine structure of the tarsal glands of the honeybee Apis mellifera L. (Hymenoptera)

Summary Tarsal glands are located in the 6th tarsomere of adult honeybee queens, workers and drones. Their structural features are not cast or sex specific. The glandular epithelium is lined by a thin endocuticular layer. A cuticular pocket is formed from a postimaginal delamination of the cuticle secreted by the glandular epithelium. The apical plasma membrane of the glandular cells shows numerous cristae and microvilli lining large crypts that communicate with the subcuticular space. Pinocytotic vesicles, multivesicular bodies and residual dense bodies are present in the apical part of the glandular cells. The RER is well developed in perinuclear and basal parts of the glandular cells, but the Golgi apparatus is a discrete organelle without secretory granules. No exocytotic secretory structures were observed. To reach the glandular pocket, the non-proteinaceous secretory product must pass across the subcuticular space, the cuticular intima, the space between the intima and the cuticular wall, and the cuticular wall of the glandular pocket.     

A histochemical study of the reproductive structures in the flatworm Dugesia leporii (Platyhelminthes, Tricladida)

Abstract. The functional morphology and the topographic distribution of tissues in the reproductive system of specimens of Dugesia leporii , an endemic Sardinian free-living planarian, are investigated. Data are provided on the nature of epithelial and glandular secretions, spermatophores, and cocoons by histochemistry, light microscopy, and scanning electron microscopy. All secreting epithelial cells produce strongly acidic sulfated glycoproteins. Glandular cells secrete strongly acidic sulfated glycoproteins or keratohyalin-like material in the penis bulb, and prekeratin-like material in atrial glands. Secretions of the bursa copulatrix may be involved in the activation of sperm while material produced by the bursa canal and oviducts probably serves to propel spermatophores or sperm and eggs. Mucous secretion of the seminal vesicle may serve to dilute and activate sperm before copulation. The viscous secrete of the ejaculatory duct and vasa deferentia may play a protective role to maintain sperm viability. Materials produced by the penis papilla and atrium probably lubricate the epithelial surface. The bilayered wall of spermatophore made of keratohyalin-like material and strongly acidic sulfated glycoproteins is produced by two gland types of the penis bulb. The bilayered shell of cocoon made of prekeratin-like and keratohyalin-like materials is secreted by both atrial glands and vitelline cells. The cocoon stalk is made of keratohyalin-like material produced by cement glands. Shell glands, producing GAG, are not involved in cocoon formation, but they may be implicated in the dilution and activation of seminal material to favor sperm movement toward the oviducts.     

Morphology of the male system and spermatophores of the arrowworm Ferosagitta hispida (Chaetognatha)

Transmission electron microscopy reveals that the somatic testicular tissues and sperm ducts are elaborations of the epithelial lining of the tail coelom. The testes consist of closely packed spermatogonia embedded between specialized lateral field cells. These cells contain few organelles and appear to function mainly as a compartment boundary. Masses of spermatogenic cells are released into the tail coelom from the anterior end of the testes. The sperm ducts, lined by simple cuboidal ciliated epithelium, collect mature spermatozoa from the tail coelom and convey them to the blindly ending seminal vesicles. The sperm ducts also modify coelomic fluid entering them along with the spermatozoa. The seminal vesicles consist of a simple glandular lining epithelium embedded in the stratified epidermis. Secretions of the lining epithelium surround the enclosed sperm mass and correspond in position to a noncellular spermatophore coat visible by light microscopy around released sperm masses. Spermatophores leave the seminal vesicles through a temporary split that forms between microfilament-containing suture cells. Maturation of spermatozoa and filling of the seminal vesicles is cyclical, occurring late each day. © 1994 Wiley-Liss, Inc.     

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.     

Fine Structure of Female Accessory Reproductive Gland in the Mealworm Beetle, Tenebrio molitor

ABSTRACT The fine structure of female accessory reproductive gland (FARG) of the adult mealworm beetle, Tenebrio molitor is studied with light and electron microscopes. The FARG is a simple tubular organ that composed of two kinds of cells-secretory epithelial cells and duct forming cells. The lumen of FARG is lined with a thin cuticle and filled with secretory materials. Each secretory epithelial cell has its peculiar end apparatus in addition to well-developed rough endoplasmic reticulum (rER), mitochondria, and secretory vesicles. They are forming basal infolding along the plasma membrane. Along the inner surface of the plasma membrane, numerous secretory vesicles are seen. The glandular secretions of the epithelial secretory cells are synthesized via rER to Golgi apparatus, and are stored in the extracellular cavity in the epithelial cell. These secretions are drained to the lumen through the end apparatus and this type of glandular secretion in the insects is type III. Histochemical reactions reveal the major component of these glandular secretions is an acid mucopolysaccharide.     

Scanning microscopy of the developing vagina in postnatal gerbils

Scanning electron microscopy of postnatally developing gerbil vagina (birth to maturity) shows that longitudinal folds form prior to transverse folds; the process of fold formation is initiated on the dorsal wall and proceeds ventrally. From days 1 to 7 postnatally, the vaginal epithelium is composed of either flat or bulging cells, depending on the vaginal region. The luminal cell surface is covered with uniform stubby microvilli and solitary cilia. Between days 9 and 20, the flat cells with distinct cell boundaries spread toward more proximal areas, leading to the formation of mixed patches of cells with flat or rounded apices. Individual elongated microvilli or tufts of forked microvilli may sprout from their surfaces. Solitary cilia gradually disappear. The transition from immature to mature vaginal epithelium starts around day 20, when individual cells recess below the level of neighboring cells. This process spreads throughout the vagina during the following days, reflecting local changes in the subsurface layers of the epithelium preparatory to exfoliation. Around day 40 the actual exfoliation of the luminal cell layer starts. By this time the surface characteristics of many of the desquamating cells have changed. In addition to microvilli, microridges are being formed. The process of exfoliation is finished by about day 60. The newly appearing cell layers now transform into typical cornified cells of the cycling vaginal epithelium.     

Sperm storage by spermatodoses in the spermatheca of Trioza alacris (Flor, 1861) hemiptera,psylloidea, triozidae: A structural and ultrastructural study

Female insects generally store sperm received during mating in specific organs of their reproductive tract, i.e., the spermathecae, which keep the sperm alive for a long time until fertilization occurs. We investigated spermatheca morphology and ultrastructure in the psylloidean insect Trioza alacris (Flor, 1861 ) in which spheroidal sperm packets that we refer to as ‘spermatodoses’ are found after mating. The ectoderm‐derived epithelium of the sac‐shaped spermatheca that has a proximal neck, consists of large secretory and flat cuticle‐forming cells. Secretory cells are characterized by a wide extracellular cavity, bordered by microvilli, in which electron‐dense secretion accumulates before discharge into the spermathecal lumen. The cuticle‐forming cells produce the cuticular intima of the organ and a peculiar specialized apical structure, through which secretion flows into the lumen. At mating, the male transfers bundles of sperm cells embedded in seminal fluid into the spermathecal neck. Sperm cells proceed towards the spermathecal sac lumen, where they are progressively compacted and surrounded with an envelope that also encloses secretions of both male and female origin. We describe the formation of these sperm containing structures and document the contribution of the female secretion to spermatodose or female‐determined spermatophore construction. We also discuss the choice of the term ‘spermatodose’ for T. alacris and suggest it be used to refer to sperm masses constructed in the female reproductive organs, at least when they involve the contribution of female secretion. © 2011 Wiley Periodicals, Inc.     

Specific criteria in Grania (Oligochaeta,Enchytraeidae)

The structure of the penial bulb and male efferent duct system of Grania species may be used in addition to setal pattern and spermathecal shape to distinguish species. Six penial bulb types are distinguished: (1) a simple, small, glandular bulb surrounding the male pore; (2) a small, glandular bulb, with a large, associated, dorso-medial gland mass; (3) a small glandular bulb, medial to the male pore, with an elongate male bursa (the aglandular sac), the vas deferens exitting directly into the invaginated male pore; (4) a glandular bulb with an aglandular sac and a small, cuticular stylet embedded in the bulb, extending from the ectal end of the vas deferens; (5) a glandular bulb and an aglandular sac with a long stylet extending from the vas deferens, through the bulb into the sac; and (6) glandular bulb reduced or absent, with or without an aglandular sac; with a long stylet and other prominent modifications, usually muscular, of the vas deferens. The details of the male duct structure were consistent within specimens grouped on the basis of setal distribution and shape and detailed spermathecal structure. Diverse male duct patterns are found within the polytypic species G. macrochaeta and G. postclitellochaeta. The positions of the spermathecal and male pores in their respective segments are distinctive for some species.     

The uterovaginal sperm host glands of the quail (Coturnix coturnix japonica)

Summary Ultrastructural and ultrahistochemical studies were performed on the uterovaginal sperm host glands of the quail (Coturnix coturnix japonica). The proximal parts of the glandular necks are lined by a pseudostratified epithelium, consisting of high columnar ciliated cells and small, irregular shaped, basal cells.The true glandular epithelium is composed only of columnar cells with microvilli on their luminal end. A characteristic luminal feature is a large lipid droplet in the perinuclear region. In the subplasmalemmal region numerous tubular profiles are seen which could represent a cellular resorption system.To evaluate the absorptive capacity of the Uterovaginal sperm host glands, tracer studies with HRP, ferritin, lanthanum and ruthenium red were undertaken. Since between 5 min and 3 h after injection no absorption could be found with the techniques mentioned, it is suggested that phagocytosis of spermatozoa by the glandular epithelium is not likely to occur.     

Morphology of defense glands of the opilionids (daddy longlegs) Leiobunum vittatum and L. flavum (Arachnida: Opiliones: Palpatores: Phalangiidae)

Opilionid defense glands consist of 0.5 × 0.9-mm sacs attached to the underside of low tubercles located on the dorsal side of the cephalothorax, posterior to the first pair of legs. Each gland opens via an elongated slit, located in the posterior floor of a crater that is situated at the summit of the tubercle. The center of the sac, called the reservoir, is lined by a cuticle consisting of epicuticle and endocuticle which is continuous through the slit with the exoskeleton. The layers of cuticle vary in thickness with different locations in the gland. A hemocoelomic (basement) membrane, 0.5–1, μ thick, forms the boundary between glandular cells and hemocoel. The gland has a nonsecretory portion consisting only of cuticle-supporting cells and a secretory portion consisting of secretory and cuticle-supporting cells. The cuticle lining the reservoir in the secretory area is broached by many cuticle-lined ductules, each of which drains an isolated intercellular space called the intercalated cistern. This in turn drains microvilli-lined canaliculi located between and extending into secretory cells. The cisterns are devoid of microvilli. Secretory cell cytoplasm contains a Golgi apparatus, many free ribosomes, rough endoplasmic reticulum (RER), two types of granules (speckled and dense), and mitochondria. Speckled granules are partially filled with fairly large particles and are found in association with the Golgi apparatus. They also surround canaliculi into which they empty. Dense granules are packed with very small particles, have a gray homogeneous appearance, and are scattered throughout the cytoplasm. Mitochondria containing matrix granules tend to scatter throughout the cytoplasm but are concentrated around canaliculi.     

Ultrastructure of sperm storage and male genital ducts in a male holocephalan, the elephant fish, Callorhynchus milii.

In chondrichthyes, the process of spermatogenesis produces a spermatocyst composed of Sertoli cells and their cohort of associated spermatozoa linearly arrayed and embedded in the apical end of the Sertoli cell. The extratesticular ducts consist of paired epididymis, ductus deferens, isthmus, and seminal vesicles. In transit through the ducts, spermatozoa undergo modification by secretions of the extratesticular ducts and associated glands, i.e., Leydig gland. In mature animals, the anterior portion of the mesonephros is specialized as the Leydig gland that connects to both the epididymis and ductus deferens and elaborates seminal fluid and matrix that contribute to the spermatophore or spermatozeugmata, depending on the species. Leydig gland epithelium is simple columnar with secretory and ciliated cells. Secretory cells have periodic acid-Schiff positive (PAS+) apical secretory granules. In the holocephalan elephant fish, Callorhynchus milii, sperm and Sertoli cell fragments enter the first major extratesticular duct, the epididymis. In the epididymis, spermatozoa are initially present as individual sperm but soon begin to laterally associate so that they are aligned head-to-head. The epididymis is a highly convoluted tubule with a small bore lumen and an epithelium consisting of scant ciliated and relatively more secretory cells. Secretory activity of both the Leydig gland and epididymis contribute to the nascent spermatophores, which begin as gel-like aggregations of secretory product in which sperm are embedded. Fully formed spermatophores occur in the ductus. The simple columnar epithelium has both ciliated and secretory cells. The spermatophore is regionalized into a PAS+ and Alcian-blue-positive (AB+) cortex and a distinctively PAS+, and less AB+ medulla. Laterally aligned sperm occupy the medulla and are surrounded by a clear zone separate from the spermatophore matrix. Grossly, the seminal vesicles are characterized by spiral partitions of the epithelium that project into the lumen, much like a spiral staircase. Each partition is staggered with respect to adjacent partitions while the aperture is eccentric. The generally nonsecretory epithelium of the seminal vesicle is simple columnar with both microvillar and ciliated cells.     

Sperm aggregations in the spermatheca of the red back salamander (Plethodon cinereus)

The spermatheca of Plethodon cinereus is a compound tubular gland that stores sperm from mating in early spring (March–April) to oviposition in summer (June–July). The seasonal variation of sperm storage in this species has previously been studied by light and transmission electron microscopy. In this paper, sperm aggregations, interaction of sperm with the spermathecal epithelium, and spermathecal secretions are studied using scanning electron microscopy. Within spermathecal tubules, relatively small groups of sperm are aligned along their entire lengths in parallel arrays. This pattern is similar to other plethdontids with complex spermathecae. Lumina of spermathecal tubules are filled with secretory material in April prior to the arrival of sperm, and after sperm appear, a coating of secretory material persists on the apices of the spermathecal epithelium. Sperm peripheral to the central luminal mass can become embedded in the secretory matrix or pushed deeper into the spermathecal epithelium. The spermathecal secretions may serve to attract and prolong the viability of sperm, but sperm that become enmeshed in the secretions or epithelium are phagocytized. Sperm and spermathecal secretions are largely absent after ovulation and in summer months, and new secretory vacuoles are formed in fall, although mating does not occur until spring.     

Fine structure of the excretory system of the deep posterior (Ebner's) salivary glands of the human tongue

Human deep posterior lingual glands (von Ebner's glands) are located beneath the circumvallate papillae. They are formed by tubuloalveolar adenomeres, intercalated ducts and excretory ducts coming together in the main excretory duct. The tubuloalveolar cells, pyramid-shaped, show large and dense secretory granules (clear cored) throughout the cytoplasm, rare basal folds and packed cisternae of rough endoplasmic reticulum (RER) at the basal pole. The columnar cells of the intercalated ducts are arranged in a monolayer. They are characterized by dense, clear-core secretory granules (mostly in the apical cytoplasm), a basal nucleus, well-developed RER and Golgi apparatus, and thin filaments distributed in supra- and perinuclear cytoplasm. Striated ducts are absent. Excretory ducts, coming together in the main duct, are lined by a bistratified epithelium. The inner layer consists of columnar cells showing bundles of tonofilaments with scarce secretory activity. The outer layer is composed of basal cells lying on the basal lamina. The main excretory duct, which opens at the bottom of the vallum, shows a stratified epithelium. The outer side is composed of 2-3 layers of malpighian cells lying on the basal lamina. The inner side consists of a single layer of cuboidal-columnar cells with dense apical granules and well-developed organelles synthesizing and condensing secretions. These cells interpolate with goblet cells, rare mitochondria-rich cells, ciliated cells and numerous small globous cells showing a clear matrix and lacking secretory granules. The cilia show a 9 + 2 microtubular structure with basal bodies provided with striated rootlets. Myoepithelial cells surround with their processes the basal portions of the secretory cells and the intercalated ducts. The conclusions concern some comparative aspects and some hypothesis on the functional role of goblet cells, ciliated cells and epithelial cells lining the different ducts, also in relation to the final secretory product.     

Functional morphology of the glandula prostatica,ejaculatory valve,and ductus ejaculatorius of the silkworm Bombyx mori

The penis of the silkmoth, Bombyx mori, consists of two parts covered with cuticle, the corpus penis and crus penis, and a third part, the radix penis, without a cuticle but surrounded by a thick sphincter. The radix penis is divisible into anterior and posterior parts. The ductus (d.) ejaculatorius passing through the penis has no secretory cells. In the anterior radix penis, the wall of the d. ejaculatorius is thin and without folds; in the posterior section, it is thick, with folds in its lumen. The glandula (g.) prostatica is divisible into anterior and posterior parts according to differences in the histological and morphological characteristics of the cells and their secretions, which contain many heterogeneous substances. In the anterior g. prostatica, secretions accumulate separately in the anterior and posterior sections before ejaculation. Unlike the posterior region, the anterior region displays a large mass(es) at the periphery of the lumen along the secretory cell layer. Judging from staining properties, the pearly body and the first layer of the spermatophore wall, which, after copulation, form in the female bursa copulatrix, seem to be derived from the secretions of the anterior and posterior regions of the g. prostatica, respectively. The secretion of the posterior g. prostatica contains initiatorin, which acts as a sperm-activating factor in the inner and outer matrices of the spermatophore. An ejaculatory valve is found between the radix penis and the g. prostatica. The opening of this valve is regulated by the surrounding sphincter, thus impeding the back-flow of secretions and seminal fluid in the radix penis and resulting in their transport outwards during ejaculation. The musculature of the d. ejaculatorius and the corpus penis promotes further transport of these secretions into the female bursa copulatrix.