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.     

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.     

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.     

Ultrastructural changes in the pars distalis of the maturing male rat

Anterior pituitary glands of male rats (2, 3, 5, 8, 12, 25, 36, 52, 56, and 62 days of age) were processed for electron microscopy. During early postnatal stages secretory cells are found in various stages of differentiation and comparatively few secretory granules are seen. Nuclei are mostly irregular, and the nucleo-cytoplasmic ratio is large. Many free ribosomes are present; the endoplasmic reticulum is generally sparse and the Golgi complex small or invisible. Cells are of variable shape, and numerous cytoplasmic processes project into large intercellular spaces. Many electron-dense cells which often contain myelinlike figures are seen. Lysosomes and lysosomal precursors are frequently found in secretory cells, predominantly in somatotrophs, of all immature glands. Mitotic figures are numerous in early stages after brith and decrease in number as the gland grows in size. A gradual increase in cytoplasmic volume with concomitant differentiation of cytoplasmic components as well as accumulation of secretory granules, accompanied by loss of myelin-like figures and decrease in the number of electron-dense cells, is observed as the animal reaches the prepuberal stage. Few lysosomes are seen in cells of mature glands. At 36 days of age all secretory cells seem to have differentiated, and morphological features as well as granule content show little change until puberty is reached. Gonadotrophs attain their characteristic morphology later than other cells. Cilia are observed in all developmental stages but are relatively infrequent in the mature gland. The described ultrastructural characteristics reflect the degree of maturation as well as the functional capacities of secretory cells at particular stages of development.     

A pair of rosette glands in the embryo and zoeal larva of an estuarine crab Sesarma haematocheir, and classification of the tegumental glands in the embryos of other crabs

A pair of rosette glands (one of the tegumental glands in crustaceans) is present at the root of the dorsal spine of the thorax in mature embryos of the estuarine crab Sesarma haematocheir. Each rosette gland is spherical, 45-50 microm in diameter. This gland consists of three types of cells: 18-20 secretory cells, one central cell, and one canal cell. The secretory cells are further classified into two types on the basis of the morphology of secretory granules. There are 17-19 a cells, and only one b cell per rosette gland. An a cell contains spherical secretory granules of 2-3 microm in diameter. The granules are filled with highly electron-dense materials near the nucleus but have lower electron-density near the central cell. The secretory granules contained in the b cell have an irregular shape and are 1-1.5 microm in diameter. The density of the materials in the granules is uniform throughout the cytoplasm. The secretory granules contained in both the a and b cells are produced by the rough endoplasmic reticulum. Materials in the granules are exocytotically discharged into the secretory apparatus inside the secretory cell, sent to the extracellular channels in the central cell, and secreted through the canal cell. The rosette gland can be distinguished from the epidermal cells 2 weeks after egg-laying and the gland matures just before hatching. Materials produced by this gland are secreted after hatching and secretion continues through five stages of zoeal larvae. These rosette glands were never found in the megalopal larva. Rosette glands are found in the embryos of Sesarma spp. and Uca spp. In other crabs, tegumental glands are also found at the same position as in the embryo of S. haematocheir, but the fine structure of their glands is largely different from that of the rosette gland. On the basis of the morphology of secretory cells (a-g cell types), the tegumental glands of a variety of crab embryos can be classified into four types, including rosette glands (type I-IV). The function of these tegumental glands is not yet known, but different types of the gland seem to reflect the phylogeny of the crabs rather than differences of habitat.     

Fine structure of cells specialized for secretion of aggregation pheromone in a nitidulid beetle Carpophilus freemani (coleoptera: Nitidulidae)

The cells that secrete the aggregation pheromone of the male nitidulid beetle Carpophilus freemani are exceptionally large and lie within the body cavity. These secretory cells share many ultrastructural features with cells of other pheromone and defense glands, but they also have several unique features. A deep invagination of the surface of each of these cells acts as the secretory surface for the pheromone. The invaginated surface is highly convoluted and surrounds a narrow cuticular ductule that is connected to the tracheal system. This surface is not covered with microvilli as the comparable surfaces are in other insect secretory cells. Each secretory cell is filled with an abundance of lipid spheres that presumably contain precursors for the pheromone. Examining cells from beetles producing different levels of pheromone showed that sizes of secretory cells are positively correlated with rates of pheromone production. Whereas secretory and ductule cells of other insect glands are usually epidermal cells, these cells of nitidulid beetles represent the first pheromone glands in which oenocytes are believed to have been recruited for pheromone production and tracheal cells have been recruited as ductules for these cells.     

Fine structural aspects of secretory processes in a pentastomid arthropod parasite in its mouse and rattlesnake hosts

The histology and development of three extensive glands in the porocephalid pentastomid Porocephalus crotali is described by light and electron microscopy, during growth of the parasite to an infective stage in the tissues of mouse; the infective stage in rattlesnake definitive hosts is also included. These glands elaborate excretory/secretory components which are channelled, via chitin-lined efferent ductules, on to the parasite cuticle. Hook and frontal glands are relatively compact, and within each gland ductules serving individual secretory lobules collect into common ducts which discharge over each of the four hooks, or at the anterior margin of the cephalothorax respectively. Subparietal gland cell lobules, composed of two large and two small secretory cells, are distributed under the cuticle and each is served by a single efferent ductule; these erupt over the entire cuticle. The large cells in subparietal glands secrete lamellate droplets which coat the cuticle with thin layers. Identical cells are found in hook and frontal glands, in addition to to three morphologically distinct types of protein secretory cell. Preliminary data on the composition and immunological properties of the various secretory products are presented.     

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.     

The development of amphids and amphidial glands in adult Syngamus trachea (Nematoda: Syngamidae)

Nematode amphids are a pair of lateral cephalic sense organs, each comprising a group of sensory endings terminating in a cuticle-lined pit. In Syngamus trachea, a parasite of birds, each amphid is surrounded by two non-nervous supporting elements, a large gland cell basally and a smaller supporting cell anteriorly. The amphidial glands display high levels of secretory activity from five to six days postinfection. Secretory material is discharged through the lumen of the sense organ onto host tissue. The ultrastructure of amphids and amphidial glands has been investigated in newly moulted, immature and mature adults to trace the development of glandular activity and its effect on amphid-amphidial gland relationships. In newly moulted adults, the glands have very low levels of secretory activity and appear to act only as supporting cells to the amphids. As secretory activity increases, the gland cell membrane surrounding the sensory endings is elaborated into a reticulum which probably forms the secretory surface. In mature adults the amphid pit is swollen and filled with secretion; the sensory endings are relegated to the periphery of the lumen. It is suggested that amphidial glands develop from typical supporting cells, but acquire a new role possibly associated with parasite attachment.     

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.     

Fine structure of the pygidial glands of Bledius mandibularis (coleoptera : staphylinidae)

The pygidial glands of B. mandibularis produce a mixture of terpenes, fatty acid derivatives, and a benzoquinone. The morphology of these glands is described with particular attention to the ultrastructure of the secretory cells and their efferent ductules. Each functional secretory unit consists of two secretory cells (cortical and medullary) both of which are associated with a common extracellular cuticular ductule. The fenestrated tip of the ductule lies in a cavity bounded by the invaginated plasma membrane of the cortical cell; within the cavity surrounded by the medullary cell, the ductule is divided into a bulb region (where a spherical mass of fine cylinders surrounds the ductule itself) and an unfenestrated switchback region. Inflated cisternae of rough endoplasmic reticulum, filled with flocculent material of low electron density, are abundant in the cortical cytoplasm, and presumably represent primary secretory product en route to the cavity of this cell. The plasma membrane bounding this cavity is much infolded, and the inner surface of this membrane is studded with fine particles. In contrast, few cisternae are inflated in the medullary cell and the corresponding infolded plasma membrane is smooth. The manner in which both cells may cooperate to produce the heterogeneous secretory product is discussed.     

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.     

Morphological, histochemical and immunohistochemical characterization of secretory production of the ciliary glands in the porcine eyelid

In addition to performing general histology and cytology of the ciliary glands of the miniature pig, we studied the localization of glycoconjugates and beta-defensins in these glands with the use of carbohydrate histochemical and immunohistochemical methods. The secretory cells of the glands were equipped with non-homogeneous secretory granules, a well-developed Golgi apparatus and rough endoplasmic reticulum. The secretory epithelium and luminal secretion of the glands contained large amounts of neutral and acidic glycoconjugates with various saccharide residues (alpha-L-Fuc, beta-D-Gal, alpha-D-GalNAc and sialic acid). The sebaceous glands and tarsal glands also exhibited positive reactions to most of the histochemical methods. Additionally, the antimicrobial peptide group of beta-defensins was demonstrated to be products of the ciliary glands, as well as the sebaceous glands and tarsal glands. The results obtained are discussed with regard to the specific function of the ciliary glandular secretions. These secretory products may be related to the moistening and general protection of the skin surface of the eyelid and ocular surface.     

The post-embryonic changes in Melipona quadrifasciata anthidioides Lep. (Hym. Apoidea). II. Development of the salivary glands system

The paper deals with the development of the salivary gland system in Melipona quadrifasciata anthidioides, which begins in the prepupal stage. The silk glands degenerate by autolysis at the end of the larval stage. Degeneration is characterized by cytoplasmic vacuolization and pycnosis of the nuclei of the secretory cells. The glandular secretory portion of degenerated silk glands separates from the excretory ducts. The salivary glands develop from the duct of the larval silk glands. The thoracic salivary glands develop from the ducts of the secretory tubules and the head salivary glands from the terminal excretory duct. The mandibular glands appear in the prepupa as invaginations of mandibular segments, and their differentiation to attain the adult configuration occurs during pupation. The hypopharyngeal glands have their origin from evaginations of the ventral anterior portion of the pharynx. A long tubule first appears with walls formed by more than one cellular layer. Then some cells separate from the lumen of the duct, staying attached to it by a cuticular channel in part intracellular. The initial duct constitutes the axial duct, in which the channel of the secretory cells opens. During the development of salivary and mandibular glands, they recapitulate primitive stages of the phylogeny of the bees. During the development of salivary glands system, mitosis accounts for only part of the growth. Most of the growth occurs by increase in size of cells rather than by cell division. In brown-eyed and pigmented pupae six days before emergence, the salivary gland system is completely developed, although not yet functioning.     

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.     

Biogenesis of secretory granules. Implications arising from the immature secretory granule in the regulated pathway of secretion

In endocrine cells the regulated secretion of hormones, peptides, enzymes and neurotransmitters into the external medium occurs when mature secretory granules fuse with the plasma membrane. Secretory granules form at the trans-Golgi network (TGN) by envelopment of the dense-core aggregate of regulated secretory proteins by a specific membrane. The secretory granules initially formed at the TGN, referred to here as immature secretory granules, are morphologically and biochemically distinct from mature secretory granules. The functional similarities and differences between the immature secretory granule and the mature secretory granule, and the events involved in the maturation of the secretory granules are briefly discussed.     

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.     

Cytodifferentiation in the accessory glands of Tenebrio molitor I. Ultrastructure of the tubular gland in the post-ecdysial adult male

The tubular accessory reproductive glands of the male mealworm beetle consist of a secretory epithelium surrounded by a thin muscular sheath. Each columnar secretory cell is divisible into three zones: basal which is adjacent to the muscle layer and contains rough endoplasmic reticulum and Golgi, intermediate, which contains endoplasmic reticulum and Golgi zones in the immature gland and is filled with secretory vesicles in the mature gland, and apical. Maturation also involves proliferation and organization of the rough endoplasmic reticulum in the basal and intermediate zone. The process appears to be complete at four days after ecdysis. Parallels with other insect glands and with the mammalian prostate are striking.     

Studies of the Phoront of Hyalophysa chattoni (Ciliophora,Apostomatida) Encysted on Grass Shrimp1

ABSTRACT. The apostomatous ciliate Hyalophysa chattoni, an ectosymbiont of the grass shrimp Palaemonetes pugio, encysts and dedifferentiates within 48 h from the migratory tomite to a phoretic stage devoid of complex ciliary fields. The presettlement crawling and pivoting of the tomite may play a role in its initial attachment to the shrimp. Metamorphosis of exuviotrophic apostomes has been previously observed to take place immediately prior to host ecdysis. The study has found that Hyalophysa's metamorphosis to the feeding stage on grass shrimp is initiated by a cue from the premolt host and begins during earlier stages of the molt cycle (D₀ and D₁). Due to the long premolt stage of the host's diecdysic molt cycle, metamorphosis is initiated well before ecdysis (over six days). Hyalophysa was able to encyst and metamorphose within 41/4 h when exposed to shrimp in a late premolt stage, indicating that the control of apostome metamorphosis is solely host-dependent.     

Cytodifferentiation in the accessory glands of Tenebrio molitor. XI. Transitional cell types during establishment of pattern

The bean-shaped accessory glands of male Tenebrio consist of a single-layered epithelium which is surrounded by a muscular coat. The epithelial layer, which produces precursors of the wall of the spermatophore, contains eight secretory cell types. Each secretory cell type is in one or more homogenous patches, and discharges granules which form one layer of the eight-layered secretory plug. Maturation begins in cell types 4, 7, and 6 on the last pupal day. A newly identified cell (type 8) in the posterolateral epithelium matures last. Cells of individual types mature in synchrony, and their secretory granules “ripen” in a sequence that is characteristic for each type. As the secretory cells of each patch mature, unusual short-lived cells appear at interfaces between patches. In some cases the secretory granules in these boundary cells have ultrastructural features which are mixtures of the definitive characteristics of granules in adjacent cell types. The transitional cell types disappear at 3–4 days after eclosion. Intermediate cell types are absent in the mature gland and boundaries between the patches are distinct. The transitional cells may form granules of intermediate structural characteristics as a dual response to cellular interaction with adjacent and previously differentiated secretory cells.