Structure of taste buds in foliate papillae of the rhesus monkey, Macaca mulatta

Taste buds in foliate papillae of the rhesus monkey were examined by electron microscopy. Three distinct cell types were identified. Type I cells were narrow elongated cells containing an oval nucleus, bundles of intermediate filaments, several Golgi bodies, and characteristic apical membrane-bounded dense granules. These cells exhibited morphological variations: some had a moderately dense cytoplasm, perinuclear free ribosomes, and flattened sacs of rough endoplasmic reticulum; others had a more lucent cytoplasm, dilated irregular rough endoplasmic reticulum, lysosome-like dense bodies, and lipid droplets. Type II cells typically contained a spherical, pale nucleus, a prominent nucleolus, supranuclear and infranuclear Golgi bodies, mitochondria with tubular cristae, and one or two centrioles. This cell type, too, showed some variation in the relative amounts of ribosomes and smooth endoplasmic reticulum, which varied inversely with each other. Type III cells were characterized by a clear apical cytoplasm essentially devoid of ribosomes and containing microtubules. In a few type III cells, the peri- and infranuclear regions contained many ribosomes and some rough endoplasmic reticulum. In most Type III cells, there were large numbers of dense and clear vesicles in the peri- and infranuclear regions; some of the vesicles were grouped in synapse-like arrangements with adjacent nerves. The morphological variations exhibited by all three cell types could be accounted for by age differences in each of the cells. This would be consistent with the notion that cell renewal occurs in each of the three cell populations.     

Fine structure of spermatogonia and primary spermatocytes in rabbits

Summary The fine structure of rabbit Spermatogonia and primary spermatocytes in meiotic prophase has been studied with different methods of preparation, including a technique for acid phosphatase activity. The spermatogonial cytoplasm is rich in free ribosomes and containes moderate amounts of vesicular, smooth-surfaced endoplasmic reticulum and mitochondria, a simple Golgi-apparatus, some micropinocytotic vesicles, and occasional multivesicular bodies, vacuoles and dense bodies with acid phosphatase activity. The large type A Spermatogonia have a prominent nucleolus and their mitochondria sometimes form clusters with a dense intermitochondrial substance, similar to that in spermatocytes.The nucleus and cytoplasm of primary spermatocytes increase markedly in volume and density during meiotic prophase. The Golgi apparatus enlarges and becomes more differentiated and finally forms small proacrosome granules. The endoplasmic reticulum produces numerous small, mainly smooth vesicles and might also be the source of a new organelle: numerous piles of narrow cisternae with opaque contents. These piles disintegrate late in prophase. The mitochondria become aggregated in clusters with dense intermitochondrial substance and their internal structure is characterized by highly dilated cristae and small particles, interpreted as mitochondrial ribosomes, in the matrix. The role of these structures in the formation of new mitochondria is discussed. The clusters of mitochondria finally disperse and their cores of dense intermitochondrial substance, possibly containing ribonucleoprotein, coalesce into a large chromatoid body similar to that in spermatids. Micropinocytosis and a few lysosomes occur in most spermatocytes. The pachytene nuclei show prominent nucleoli and a distinct sex vesicle without any synaptinemal complex.The importance for spermatid differentiation of some events taking place in the cytoplasm of primary spermatocytes is emphasized.Financial support for this study was received from the Swedish Medical Research Council.     

长吻鮠精巢发育的分期及精子的发生和形成

长吻鮠精巢的发育分为精原细胞增殖期、精母细胞生长期、精母细胞成熟期、精子细胞出现期,精子完全成熟期和精子退化吸收期。精巢的后1/3不产生也不贮存精子,精子的发生和形成经过精原细胞、精母细胞、精子细胞到精子的一系列过程。精原细胞有两种类型。精子无顶体,有中心粒帽,中片长,核凹窝和线粒体发达,鞭毛具侧鳍。     

Etude ultrastructurale des spermatogonies et de la croissance spermatocytaire chez Lithobius forficatus L. (Myriapode Chilopode)

Résumé L'étude ultrastructurale des spermatogonies et des spermatocytes a été envisagée chez Lithobius forficatus L. (Myriapode Chilopode). Les spermatogonies présentent un noyau à chromatine dispersée dont le nucléole est condensé. Leur cytoplasme renferme des ribosomes, libres ou associés en polysomes; les mitochondries et les dictyosomes sont peu abondants.Au cours de la croissance spermatocytaire, le noyau et le cytoplasme augmentent considérablement de volume. La dispersion de la chromatine est plus importante et le nucléole présente de nombreuses figures de bourgeonnement. Les ribosomes, les mitochondries, les dictyosomes et le reticulum endoplasmique sont très abondants. De nouvelles formations sont observables: 1) des amas de saccules réticulaires concentriques, délimitant une zone cytoplasmique; 2) des empilements lamellaires ayant leur origine soit dans le reticulum banal, soit dans les amas concentriques.L'activité synthétique se manifeste lors de la croissance spermatocytaire par: 1) d'abondants échanges nucléo-cytoplasmiques; 2) une augmentation du nombre de mitochondries et des ribosomes; 3) une intense activité golgienne. Elle est vraisemblablement en rapport avec le gigantisme spermatocytaire observé chez les Myriapodes Chilopodes.

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Ultrastructural study of spermatogonia and spermatocyte growth of Lithobius forficatus L. (Myriapoda, Chilopoda)

Summary The ultrastructure of spermatogonia and spermatocytes of Lithobius was studied. The spermatogonia show a nucleus with dispersed chromatin and a compact nucleolus. Their cytoplasm contains free ribosomes and polysomes; mitochondria and dictyosomes are not abundant.During spermatocyte growth, nucleus and cytoplasm increase markedly in volume. The dispersion of the chromatin is more pronounced; the nucleolus shows frequent budding. Ribosomes, mitochondria, dictyosomes and endoplasmic reticulum are very numerous. New structures are (1) accumulations of concentric reticular saccules with a cytoplasmic center, (2) piles of narrow cisternae originating from the regular endoplasmic reticulum or from concentric saccules.The synthetic activity during spermatocyte growth manifests itself by (1) numerous signs of nucleo-cytoplasmic exchanges, (2) an increase in the number of mitochondria and ribosomes, (3) a high activity of the Golgi apparatus. This is probably related to the gigantism of spermatocytes in Chilopoda.

     

Androgenic gland cell structure and spermatogenesis during the molt cycle and correlation to morphotypic differentiation in the giant freshwater prawn, Macrobrachium rosenbergii

The freshwater prawn Macrobrachium rosenbergii shows three male morphotypes: blue-claw males (final stage having high mating activity), orange-claw males (transitional stage showing rapid somatic growth), and small males (primary stage showing sneak copulation). This morphotypic differentiation is considered to be controlled by androgenic gland hormone, which is probably a peptide hormone. However, its physiological roles are not fully understood. In the present study, we examined the correlation of androgenic gland cell structure to spermatogenic activity and morphotypic differentiation histologically in M. rosenbergii. spermatogenic activity showed close correlation to the molt cycle in orange-claw males and small males. spermatogonia increased in number in the late premolt stage, becoming spermatocytes in the postmolt stage, and spermatocytes differentiated into spermatozoa in the intermolt and early premolt stages. Ultrastructure of the androgenic gland was additionally compared among the molt stages, but, distinct histological changes were not observed in relation to spermatogenesis during the molt cycle. On the other hand, among the three morphotypes, the androgenic gland was largest in the blue-claw males, containing developed rough endoplasmic reticulum in the cytoplasm. These results suggest that, during spermatogenesis which is related to the molt cycle, the androgenic gland hormone is at rather constant levels and plays a role in maintaining spermatogenesis rather than directly regulating the onset of a specific spermatogenesis stage and that, during the morphotypic differentiation, the androgenic gland is most active in the blue-claw males and plays a role in regulating the observed high mating activity in M. rosenbergii.     

The fine structure of spermatogenesis in Hymenolepis diminuta (Cestoda) with a description of the mature spermatozoon

The processes of spermatogenesis and spermiogenesis in Hymenolepis diminuta were studied by electron microscopy using improved preparative techniques. Spermatogonia (Type A) are characterized by nuclei 3.79 (+/- 0.17) micrometer in diameter, dense cytoplasm packed with free ribosomes and aggregates of mitochondria. After mitoses, certain spermatogonia (Type B) assume syncytial rosettes containing eight nuclei. Primary spermatocytes maintain the rosette syncytium and have large nuclei (4.28 +/- 0.24 micrometer in diameter), smooth endoplasmic reticulum, and polysomes. The secondary spermatocyte is short-lived and is characterized by nuclei (2.0 +/- 0.11 micrometer in diai (2.0 +/- 0.11 micrometer in diameter) and perinuclear membranous lamellae. The syncytial spermatid cluster contains avoid nuclei which condense and elongate to a final diameter of 0.22 +/- 0.04 micrometer. Once elongated, these nuclei become delimited from the syncytium by invaginations of the plasma membrane. During delimitation, cortical peripheral microtubules arise beneath the spermatozoon plasmalemma and a 9 + 1 axoneme extends the length of the mature lance-shaped spermatozoon.     

Fine structure of the early stages of spermatogenesis in the Pacific oyster, Crassostrea gigas (Mollusca, Bivalvia)

The aim of this study is to describe the early stages of spermatogenesis of the Pacific oyster Crassostrea gigas using both light and electron microscopy. The gonad is formed by gonadal tubules invaginated in a connective tissue constituting a storage tissue. Myoepithelial cells surround each gonadal tubule and are associated with an acellular matrix delimiting the outer part of the tubule, the inner part is composed by intragonadal somatic cells associated with germinal lineage. Two types of spermatogonia are identified, where type I spermatogonia (Spg I) are large, scarce and pale cells leaned against the base of the tubule (nuclear diameter: 5.5+/-0.5 microm). Type II spermatogonia (Spg II) are clustered and dark cells which appear smaller than type I (nuclear diameter: 4.3+/-0.3 microm). The aspect of nuage-like material in cytoplasm is described from pale spermatogonia to primary spermatocytes (nuclear diameter: pachytene 3.6+/-0.3 microm, diplotene 3.4+/-0.3 microm), while no structure related to a chromatoid body was observed in oyster spermatocytes and spermatids.     

Spermatogenesis in the inseminating African butterflyfish Pantodon buchholzi (Teleostei: Osteoglossiformes: Pantodontidae) with the revision of residual bodies formation

The aim of this study was to analyse spermatogenesis in the African butterflyfish, Pantodon buchholzi, using transmission electron microscopy and scanning electron microscopy. P. buchholzi is the most basal teleost that exhibits insemination and produces a highly complex introsperm with the most elongate midpiece known in teleost fishes. Their early stages (spermatogonia and spermatocytes) do not differ greatly from those of other fishes, with the exception of Golgi apparatus degradation appearing as spindle-shaped bodies (SSBs). In round, early spermatids, the development of the flagellum begins after the migration of the centriolar complex towards the nucleus. Later, the elongation of the midpiece coincides with the displacement of the mitochondria and their fusion to produce nine mitochondrial derivatives (MDs). In these spermatids, the nucleus is situated laterally to the midpiece, with condensing chromatin in the centre of the nucleus. Within the midpiece, the flagellum is located within a cytoplasmic canal and is surrounded by a cytoplasmic sleeve containing fibres, MDs and a great amount of cytoplasm located on one side. During the next phase, nuclear rotation, the highly condensed chromatin is displaced to a position above the centriolar apparatus, whereas chromatin-free nucleoplasm is transferred to the cytoplasm. Later, this nucleoplasm, still surrounded by the nuclear membrane, is eliminated into the cyst lumen as the nucleoplasmic packet. Within the highly elongate spermatids, other excess organelles (SSBs, endoplasmic reticulum and mitochondria) are eliminated as residual bodies (RBs). Fully developed spermatozoa, which contain conical-shaped nuclei, eventually coalesce to form unencapsulated sperm packets (spermatozeugmata) that are surrounded by RBs at the level of the extremely elongate midpieces. Later, RBs are removed at the periphery of the cyst by means of phagocytosis by Sertoli cells.     

The testicular cycle of Gambusia affinis holbrooki (Teleostei: Poeciliidae)

Fifteen male mosquito fish ( Gambusia affinis holbrooki ) were collected in 1989 on the 15th of each month to perform a quantitative histologic study of the annual testicular cycle including a calculation of the gonadosomatic index, testicular volume, and the total volume per testis occupied by each germ cell type. The cycle comprises two periods: spermatogenesis and quiescence. The spermatogenic period begins in April with the development of primary spermatogonia into secondary spermatogonia, spermatocytes and round spermatids. In May, the first spermatogenic wave is completed and the testicular volume begins to increase up to June when the maximum testicular volume and gonadosomatic index are reached. Germ cell proliferation with successive spermatogenetic waves continues until August. In September germ cell proliferation ceases and neither secondary spermatogonia nor spermatocytes are observed. However, spermiogenesis continues until October. In November, spermiogenesis has stopped and the testis enters the quiescent period up to April. During this period only primary spermatogonia and spermatozoa are present in the testis. In addition, a few spermatids whose spermiogenesis was arrested in November are observed. Testicular release of spermatozoa is continuous during the entire spermatogenesis period. The spermatozoa formed at the end of this period (September-October) remain in the testis during the quiescent period and are released at the beginning of the next spermatogenesis period in April. Developed Leydig cells appear all year long in the testicular interstitium, mainly around both efferent ducts and the testicular tubule sections showing S₄ spermatids.     

Ultrastructure of the testis in Synbranchus marmoratus (Teleostei,Synbranchidae): the germinal compartment

Synbranchus marmoratus, is a protogynic diandric species in which two types of males, primary and secondary, are found. In both types, the germinal compartment in the testes is of the unrestricted lobular type, but in secondary (sex reversed females) males the lobules develop within the former ovarian lamellae. In the present study, the germinal compartment was examined in both types of males using light microscopy as well as scanning and transmission electron microscopy. Germinal compartment is limited by a basement membrane and contains Sertoli and germ cells. During maturation, processes of Sertoli cells form the borders of spermatocysts containing isogenic germ cells. Characteristically, type A and type B spermatogonia have a single nucleolus and grouped mitochondria associated with dense bodies or nuage. Type B spermatogonia, spermatocytes and spermatids are joined by cytoplasmatic bridges and are confined within spermatocysts. Secondary spermatocytes are difficult to find, indicating that this stage is of short duration. Biflagellated spermatozoa have a rounded head, no acrosome, and possess a midpiece consisting of two basal bodies, each of which produces a flagellum with a typical 9+2 microtubular composition. No associations occur between sperm and Sertoli cells. There were no differences between spermatogenesis in primary and secondary males in this protogynic, diandric fish.     

Ultrastructural localization of phosphatases in the testes of the domestic fowl: Acid phosphatase and thiamine pyrophosphatase

Summary ACPase and TPPase activity has been examined in the germinal epithelium of the testes in the domestic fowl. ACPase activity in spermatogonia and spermatocytes was confined to the Golgi complex. In spermatids ACPase activity was seen in the endoplasmic reticulum and nuclear envelope in the phase I and especially in the phase II (the elongating phase). This activity gradually decreased during the next phase III, and had disappeared in the final phase IV. The membrane body showed ACPase reaction in the small peripheral vacuoles and cisternal structures surrounding large central vacuoles. ACPase was also present in vesicles surrounding the developing tail. Late spermatids showed an abundance of autophagic vacuoles which had a complex array of ACPase positive delimiting membranes. In Sertoli cells ACPase activity was predominant in the lysosomes. TPPase activity was seen in the cisternae of the Golgi complex in spermatogonia and spermatocytes. In spermatids activity was present in the endoplasmic reticulum during the phase II, but it is lost in later stages. The smaller vacuoles and cisternal structures in the membrane body also showed reaction products. According to the present results it is thought likely that the smaller vacuoles and cisternal structures of the membrane body are of endoplasmic reticulum origin. The autophagic vacuoles in spermatids and the lysosomes of Sertoli cells are considered responsible for the degradation of residual bodies cast off by spermatids.     

毛竹茎纤维次生壁形成过程的超微结构观察

利用透射电镜观察了毛竹（Ｐｈｙｌｌｏｓｔａｃｈｙｓ ｐｕｂｅｓｃｅｎｓ Ｍａｚｅｌ）茎纤维发育过程中次生壁的形成过程。纤维发育早期,细胞具有较大的细胞核和核仁;细胞质浓稠,具有核糖体、线粒体和高尔基体等细胞器。随着纤维次生壁的形成,细胞壁加厚,细胞质变得稀薄,内质网和高尔基体的数量明显增加,并且两者共同参与了运输小泡的形成;在质膜内侧可观察到大量周质微管分布。随着次生壁的进一步加厚及木质化,细胞壁     

Transition of basic protein during spermatogenesis of <Emphasis Type="Italic">Fenneropenaeus chinensis</Emphasis> (Osbeck, 1765)

According to the ultrastructural characteristic observation of the developing male germ cells, spermatogenesis of the crustacean shrimp, Fenneropenaeus chinensis, is classified into spermatogonia, primary spermatocytes, secondary spermatocyte, four stages of spermatids, and mature sperm. The basic protein transition during its spermatogenesis is studied by transmission electron microscopy of ammoniacal silver reaction and immunoelectron microscopical distribution of acetylated histone H4. The results show that basic protein synthesized in cytoplasm of spermatogonia is transferred into the nucleus with deposition on new duplicated DNA. In the spermatocyte stage, some nuclear basic protein combined with RNP is transferred into the cytoplasm and is involved in forming the cytoplasmic vesicle clumps. In the early spermatid, most of the basic protein synthesized in the new spermatid cytoplasm is transferred into the nucleus, and the chromatin condensed gradually, and the rest is shifted into the pre-acrosomal vacuole. In the middle spermatid, the nuclear basic protein linked with DNA is acetylated and transferred into the proacrosomal vacuole and assembled into the acrosomal blastema. At the late spermatid, almost all of the basic protein in the nucleus has been removed into the acrosome. During the stage from late spermatid to mature sperm, some de novo basic proteins synthesized in the cytoplasm belt transfer into the nucleus without a membrane and almost all deposit in the periphery to form a supercoating. The remnant histone H4 accompanied by chromatin fibers is acetylated in the center of the nucleus, leading to relaxed DNA and activated genes making the nucleus non-condensed.     

Spermatogenesis and related plasma androgen levels in Atlantic halibut (Hippoglossus hippoglossus L.)

Spermatogenesis in male Atlantic halibut (Hippoglossus hippoglossus L.) was investigated by sampling blood plasma and testicular tissue from 15-39-month-old fish. The experiment covered a period in which all fish reached puberty and completed sexual maturation at least once. The germinal compartment in Atlantic halibut testis appears to be organized in branching lobules of the unrestricted spermatogonial type, because spermatocysts with spermatogonia were found throughout the testis. Spermatogenesis was characterized histologically, and staged according to the most advanced type of germ cell present: spermatogonia (Stage I), spermatogonia and spermatocytes (Stage II), spermatogonia, spermatocytes and spermatids (Stage III), spermatogonia, spermatocytes, spermatids and spermatozoa (Stage IV), and regressing testis (Stage V). Three phases could be distinguished: first, an initial phase with low levels of circulating testosterone (T; quantified by RIA) and 11-ketotestosterone (11-KT; quantified by ELISA), spermatogonial proliferation, and subsequently the initiation of meiosis marked by the formation of spermatocytes (Stage I and II). Secondly, a phase with increasing T and 11-KT levels and with haploid germ cells including spermatozoa present in the testis (Stage III and IV). Thirdly, a phase with low T and 11-KT levels and a regressing testis with Sertoli cells displaying signs of phagocytotic activity (Stage V). Circulating levels of 11-KT were at least four-fold higher than those of T during all stages of spermatogenesis. Increasing plasma levels of T and 11-KT were associated with increasing testicular mass throughout the reproductive cycle. The absolute level of, or the relation between, testis growth and circulating androgens were not significantly different in first time spawners compared to fish that underwent their second spawning season. These results provide reference levels for Atlantic halibut spermatogenesis.     

Ultrastructural Study of Secondary Wall Formation in the Stem Fiber of Phyllostachys pubescens

Ultrastructural changes in secondary wall formation of Phyllostachys pubescens Mazel fiber were investigated with transmission electron microscopy. Fiber developed initially with the elongation of cells containing ribosomes, mitochondria and Golgi bodies in the dense cytoplasm. During the wall thickening, the number of rough endoplasmic reticulum and Golgi bodies increased apparently. There were two kinds of Golgi vesicles, together with the ones from endoplasmic reticulum formed transport vesicles. Many microtubules were arranged parallel to the long axis of the cell adjacent to the plasmalemma. Along with the further development of fiber, polylamellate structure of the secondary wall appeared, with concurrent agglutination of chromatin in the nucleus, swelling and disintegration of organelles, while cortical microtubules were still arranged neatly against the inner side of plasmalemma. Lomasomes could be observed between the wall and plasmalemma. The results indicated that the organelles, such as Golgi bodies together with small vesicles, rough endoplasmic reticulum and lomasomes, played the key role in the thickening and lignification of the secondary wall of bamboo fiber, though cortical microtubules were correlative with the process as well.     

Histology of salmonid testes during maturation

The commonly applied classification systems of fish gonad maturity divide the maturation process into certain stages. However, the scales do not entirely reflect the continuity of the maturation process. Based on light microscope observations, the paper describes a comprehensive pattern of testicular transformations during maturation. The study was carried out on precocious underyearling and 1-year-old males of sea trout (Salmo trutta m. trutta L.), 1-year-old males of salmon (Salmo salar L.), and males of brown trout (Salmo trutta m. fario L.) aged from 7 months to 4 years. A total of 821 gonads collected during all seasons of the year were examined. The fish were fixed in Bouin's fluid. Histological slides of the mid-part of the gonad were made using the standard paraffin technique. The 3-6 microm sections were stained with Heidenhain haematoxylin. Histological changes of testes during maturation were similar in the three species studied. Immature and resting gonads contained type A spermatogonia in lobules only. The appearance of cystic structures containing type B spermatogonia in the lobules signalled the beginning of the sexual cycle in male gonads. Type B spermatogonia underwent synchronous mitotic divisions resulting in an increase in the total number of spermatogonia. As the spermatogenesis continued, the gonads showed a gradual increase in the number of cysts containing cells at all the spermatogenetic stages: type B spermatogonia, primary and secondary spermatocytes, spermatids, and spermatozoa. The well-formed spermatozoa were released to the lobule lumen once the Sertoli cells and spermatozoa connections broke up and the cyst disappeared. This was a continuous process observed throughout the spawning season. The spermatozoa were moved to the efferent duct. While some of the germ cells were completing spermatogenesis, the lobules contained less and less cysts with type B spermatogonia, primary and secondary spermatocytes, and spermatids; eventually all the cells completed spermatogenesis. At the end of maturation, vacuoles, up to 18.9 microm in final diameter (brown trout), appeared in the Sertoli cells. The vacuoles were visible in the lobule wall epithelium for a prolonged period of time. In most salmonid individuals examined, the reproductive cycles were observed to overlap. In some fish, the preparation for another cycle began very early, i.e., at the and of preceding spermatogenesis, which had not been observed before. Gonad maturation in some males was incomplete.     

Small granule-containing cells of the central nervous system of the bivalve mollusk Patinopectin yessoensis (Jay)

In the CNS of the Patinopecten yessoensis (Jay) two types of cells have been revealed. The I type cells are typical unipolar neurons with a developed granular endoplasmic reticulum and Golgi compex, with a nucleus containing small amount of chromatin. They possess elementary peptidergic granules. The II type cells have in their cytoplasm and processes a large amount of electron-opague granules, specific for adrenergic systems. The nucleus is rich in clustered chromatin, the granular endoplasmic reticulum is poorly developed, cytosomes are absent. According to their ultrastructural organization the latter correspond to small granular cells of the mammalian autonomic nervous system.     

Fine structure of the dorsal epithelium of the tongue of the freshwater turtle,Geoclemys reevesii (Chelonia,Emydinae)

Scanning electron microscopy shows that lingual papillae occur all over the dorsal surface of the tongue of the freshwater turtle, Geoclemys reevesii. The surface of each papilla is composed of compactly distributed hemispherical bulges, each composed of a single cell. Microvilli are widely distributed over the surface of cells. Histological examination reveals that the connective tissue penetrates deep into the center of papillae and that the epithelium is stratified columnar. Under the transmission electron microscope, the cells of the basal and the deep intermediate layers of the epithelium appear rounded. A large nucleus lies in the central area of each cell. The cytoplasm contains mitochondria, endoplasmic reticulum and free ribosomes. The cell membrane form numerous processes. The shallow intermediate layer contains two types of cell. The cytoplasm of the first has numerous fine granules, in addition to mitochondria, ribosomes, and endoplasmic reticulum. The other type of cell contains highly electron-dense granules. The surface layer shows two cell types. One type consists of typical mucous cells. The other type of cell contains fine, electron-lucent granules. The latter cells lie on the free-surface side, covering the mucous cells, and have microvilli on their free surfaces.