Comparing the secretory pathway in honeybee venom and hypopharyngeal glands

We provide insights into the secretory pathway of arthropod gland systems by comparing the royal jelly-producing hypopharyngeal glands and the venom-producing glands of the honeybee, Apis mellifera. These glands have different functions and different product release characteristics, but both belong to the class 3 types of insect glands, each being composed of two cells, a secretory cell and a microduct-forming cell. The hypopharyngeal secretory cells possess an extremely elongate tubular invagination that is filled with a cuticular structure, the end-apparatus, anchored against the cell membrane by a conspicuous series of actin rings. In contrast, venom glands have no actin rings, but instead have an actin-rich brush border surrounding the comparatively short and narrow end-apparatus. We relate these cytoskeletal differences to the production system and utilisation of secretions; venom is stored in a reservoir whereas royal jelly and enzymes are produced on demand. Fluorescence-based characterisation of the actin cytoskeleton combined with scanning electron microscopy of the end-apparatus allows for detailed characterisation of the point of secretion release in insect class 3 glands.     

The fine structure of the female reproductive system of Zorotypus caudelli Karny (Zoraptera)

The general structure of the female genital system of Zorotypus caudelli is described. The ovarioles are of the panoistic type. Due to the reduction of the envelope (tunica externa) the ovarioles are in direct contact with the hemolymph like in some other insect groups, Plecoptera included. The calices are much larger in Z. caudelli then in Zorotypus hubbardi and their epithelial cells produce large amounts of secretions, probably protecting the surface of the eggs deposited on the substrate. Eggs taken from the calyx bear a series of long fringes, which are missing in the eggs found in the ovariole, and in other zorapteran species. The long sperm of Z. caudelli and the long spermathecal duct are likely related to a sexual isolating mechanism (cryptic female choice), impeding female re-mating. The apical receptacle and the spermathecal duct - both of ectodermal origin - consist of three cell types. In addition to the cells beneath the cuticle lining the lumen, two other cell types are visible: secretory and canal cells. The cytoplasm of the former is rich in rough endoplasmic reticulum cisterns and Golgi complexes, which produce numerous discrete dense secretory bodies. These products are released into the receiving canal crossing the extracellular cavity of secretory cells, extending over a series of long microvilli. The secretion is transported towards the lumen of the apical receptacle of the spermatheca or to that of the spermathecal duct by a connecting canal formed by the canal cells. It is enriched by material produced by the slender canal cells. Before mating, the sperm cells are enveloped by a thick glycocalyx produced at the level of the male accessory glands, but it is absent when they have reached the apical receptacle, and also in the spermathecal duct lumen. It is likely removed by secretions of the spermatheca. The eggs are fertilized at the level of the common oviduct where the spermathecal duct opens. Two micropyles at the dorsal side of the equator level possibly facilitate fertilization. The presence of these two micropyles is a presumably derived feature shared with Phasmatodea. The fine structure of the female reproductive system of Z. caudelli does not allow to assess the phylogenetic position at the present stage of knowledge. The enlarged calyx and the temporary presence of long fringes on the eggs are potential autapomorphies of Z. caudelli or may indicate relationships with other Zorotypus species.     

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.     

Ultrastructural and functional aspects of the spermatheca of the African Migratory Locust Locusta migratoria migratorioides (Reiche and Fairmaire) (Orthoptera: Acrididae)

The ultrastructure of the spermathecal epithelium of the African Migratory Locust Locusta migratoria migratorioides R. & F. (Orthoptera: Acrididae) was investigated with the aid of transmission and scanning electron microscopic methods. The unpaired spermatheca can be subdivided into a multiple coiled tube and a terminal bulb region with vestibule, small apical and extensive pre-apical diverticulum. The wall of the spermatheca consists of a chitin intima, a layer of epithelial cells with a distinct apical microvilli border and a layer of gland cells, whereby slender projections of the epithelial cells extend between the gland cells. Through extensive folding, the basal plasma membrane of the gland and epithelial cells form a huge labyrinth, which is bounded by a basal lamina. Extending into the above mentioned projections there are bundles of parallel-arrayed microtubules, which run perpendicular to the microvilli border of the epithelial cell. They end in the base region of the microvilli and in the basal labyrinth on hemidesmosomes and serve to provide a mechanically stressable anchorage for the epithelium. The gland cells show structures typical for the production of export proteins: ribosomes, rER, dictyosomes, as well as vesicles of different size and electron-density. Every gland cell contains an extracellular cavity, arising through invagination, which is coated with a microvilli border. Over an end-apparatus and a ductule joining onto it (also with chitin intima) the lumen of the extracellular cavity is connected with the spermathecal lumen. The release of secretions and other substances from the epithelium into the spermatheca lumen is as possible as the uptake of substances from the latter into the epithelium. Regional differences in the fine structure of the cuticular intima, epithelial and gland cells point to different functions of the epithelium in these regions.     

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.     

Sperm storage in Spermathecae of the great lamper eel,Amphiuma tridactylum (Caudata: Amphiumidae)

The spermathecae of ten female Amphiuma tridactylum were examined by light and electron microscopy during the presumed mating and ovipository seasons (March–August) in Louisiana. Spermathecae were simple tubuloalveolar glands in the dorsal wall of the cloaca. Six of the ten specimens were vitellogenic, and all of these specimens contained sperm in their spermathecae and had secretory activity in the spermathecal epithelium. Two nonvitellogenic females also had sperm in their spermathecae and active epithelial cells, whereas the other nonvitellogenic females lacked stored sperm and secretory activity in the spermathecae. In specimens storing sperm from March–May, the sperm were normal in cytology, and secretory vacuoles were contained within the epithelium. In the August sample, however, evidence of sperm degradation was present, and secretory material had been released into the lumen by an apocrine process. We therefore hypothesize that the spermathecal secretions function in sperm degeneration. © 1996 Wiley-Liss, Inc.     

宽翅曲背蝗受精囊形态与超微结构观察

[目的]明确宽翅曲背蝗Pararcyptera microptera meridionalis雌虫受精囊的形态、组织结构与超微结构,为更好地认识昆虫受精囊的功能提供依据.[方法]本研究以宽翅曲背蝗已交配雌成虫为实验材料,利用光学显微镜和透射电子显微镜观察其受精囊的形态、组织结构和超微结构.[结果]宽翅曲背蝗受精囊由一个端囊和一条长的受精囊管组成,端囊用于储存精子.端囊和受精囊管有相似的组织学结构,由外到内依次为肌肉层、基膜、上皮层及表皮内膜.上皮层含上皮细胞、腺细胞和导管细胞3种细胞类型.腺细胞具有一个被有微绒毛的细胞外腔.腺细胞的分泌物经细胞外腔通过分泌导管进入到受精囊腔.分泌导管由导管细胞形成.[结论]在宽翅曲背蝗受精囊的端囊和受精囊管上,内膜和腺细胞的细胞外腔结构均存在差异,由此推测,端囊和受精囊管的功能存在一定差异.上皮细胞的超微结构特点显示上皮细胞具有支持、分泌和吸收的功能.     

Fine structure of the spermatheca and of the accessory glands in Orchesella villosa (Collembola, Hexapoda)

The spermatheca and the accessory glands of the collembolan Orchesella villosa are described for the first time. Both organs exhibit ultrastructural differences, according to the time of the intermolt in which the specimens were observed. A thick cuticular layer lines the epithelial cells of the accessory glands. In the reproductive phase, they are involved in secretory activity; a moderately dense secretion found in the apical cell region opens into the gland lumen. Cells with an extracellular cistern are intermingled with the secretory cells. These cells could be involved in fluid secretion, with the secretory product opening into the cistern which is filled with an electron-transparent material. After the reproductive phase, the gland lumen becomes filled with a dense secretion. The accessory gland secretion may play a protective role towards the eggs. The spermatheca is located between the accessory glands; its epithelium is lined by a thin cuticle forming spine-like projections into the lumen and consists of cells provided with an extracellular cistern. Secretory cells, similar to those seen in the accessory glands, are missing. Cells with a cistern could be involved in the production of a fluid secretion determining sperm unrolling and sperm motility.     

Fine structure of the integumental glands of a termite soldier

The ultrastructure of some integumental glands occurring in the head, thorax and abdomen of K. flavicollis soldiers is described. The secretory units consist of two cells, the canal cell and the secretory cell (this latter filled with secretion granules). A cylindrical and distorted extracellular space, or reservoir, with an irregular outline is lined by short microvilli. The end-apparatus is made up of small overlapping cuticular laminae which in section resemble small wavy rods. The ample distribution of the units has led the authors to consider them dermal glands. Scanning electron micrographs confirm that the glands' activity consists in the secretion of material which then spreads over the surface of the integument. The dissimilar appearance of the secretion granules present in glands of different soldiers suggests that the electron-lucid granules and the granules with fibrils are two completely different secretions at different ages of the animal. The authors do not therefore rule out the hypothesis that these integumental glands may later produce or release pheromones.     

The development of the female accessory gland in the insect Rhodnius prolixus

The specialized cell types and two distinct regions of the adult Rhodnius prolixus cement gland develop from a simple pseudostratified epithelial tube during the 20–22 days of the fifth stadium. Feeding initiates the first phase, proliferation. Cells round up and divide tangentially to the lumen. Following the proliferation phase, differentiative mitoses occur and differentiation, resulting in secretory units (consisting of a ductule, gland cell and cuticular lining), ensues in the distal region. Ductule morphogenesis occurs without pseudocilia, thus differing from other insect glands. The complex changes in cell shape and interaction occur during development of the secretory unit. The secretory cell and end-apparatus develop from a double cell unit at the base of elongating ductules. The inner cell produces a complex end-apparatus of epicuticle that mirrors the microvillar pattern and then it degenerates. The ductules are lined by cuticulin and inner epicuticle while the central gland lumen has a layer of endocuticle as well. The epithelium of the proximal region remains simple producing the thick corrugated cuticle characteristic of the adult secretory duct. The mesodermal covering forms a thick longitudinal striated muscle layer that adheres to the epithelium via desmosomes.     

Venom and Dufour's glands of the emerald cockroach wasp Ampulex compressa (Insecta,Hymenoptera, Sphecidae): Structural and biochemical aspects

The digger wasp species Ampulex compressa produces its venom in two branched gland tubules. They terminate in a short common duct, which is bifurcated at its proximal end. One leg is linked with the venom reservoir, the other one extends to the ductus venatus. Each venom gland tubule possesses, over its entire length, a cuticle-lined central duct. Around this duct densely packed class 3 gland units each composed of a secretory cell and a canal cell are arranged. The position of their nuclei was demonstrated by DAPI staining. The brush border of the secretory cells surrounds the coiled end-apparatus. Venom is stored in a bladder like reservoir, which is surrounded by a thin reticulated layer of muscle fibres. The reservoir as a whole is lined with class 3 gland units. The tubiform Dufour's gland has a length of about 350 μm (∅ 125 μm) only and is surrounded by a network of pronounced striated muscle fibres. The glandular epithelium is mono-layered belonging to the class 1 type of insect epidermal glands. The gland cells are characterized by conspicuous lipid vesicles. Secretion of material via the gland cuticle into the gland lumen is apparent. Analysis of the polypeptide composition demonstrated that the free gland tubules and the venom reservoir contain numerous proteins ranging from 3.4 to 200 kDa. The polypeptide composition of the Dufour's gland is completely different and contains no lectin-binding glycoproteins, whereas a dominant component of the venom droplets is a glycoprotein of about 80 kDa. Comparison of the venom reservoir contents with the polypeptide pattern of venom droplets revealed that all of the major proteinaceous constituents are secreted. The secreted venom contains exclusively proteins present in the soluble contents of the venom gland. The most abundant compound class in the Dufour's gland consisted of n-alkanes followed by monomethyl-branched alkanes and alkadienes. Heptacosane was the most abundant n-alkane. Furthermore, a single volatile compound, 2-methylpentan-3-one, was identified in various concentrations in the lipid extract of the Dufour's gland.     

Spermathecal cytology of Ambystoma opacum (Amphibia: Ambystomatidae) and the phylogeny of sperm storage organs in female salamanders

Sperm storage glands, spermathecae, were examined from mated female Ambystoma opacum during the breeding season. No differences occur in the spermathecal ultrastructure of individuals sacrificed prior to oviposition and those sacrificed within 3 days of removal from tended clutches of recently oviposited eggs. The simple tubuloalveolar glands produce two types of secretory vacuoles. Apical secretory vacuoles contain glycosaminoglycans for export into the lumen to bathe stored sperm, perhaps providing the chemical/osmotic environment necessary for sperm quiescence. The other type of secretory vacuole contains an unsaturated lipid that is produced for export into the connective tissue surrounding the spermathecae. The role of this secretion may involve the contraction of myoepithelial cells, resulting in sperm expulsion. Some sperm undergo degradation in the spermathecal epithelium, and an interepithelial leukocyte was observed in one specimen. Apical secretory vacuoles and sperm are absent from the spermathecae of a specimen sacrificed 62 days after removal from a tended egg clutch. This is the first report on the spermathecal cytology of a salamander from the Ambystomatidae, and comparisons with salamanders from other families provide a morphological basis for considering spermathecae polyphyletic within the Caudata. © 1993 Wiley-Liss, Inc.     

Honey bee males and queens use glandular secretions to enhance sperm viability before and after storage

Internal fertilization requires live sperm to be transferred from male to female before egg fertilization. Both males and females assist the insemination process by providing sperm with glandular secretions, which have been inferred to contain subsets of proteins that maintain sperm viability. Here we show that in the honeybee (Apis mellifera) secretions of the male accessory glands, the major contributors towards seminal fluid, enhance sperm survival. We further demonstrate that the protein fraction of the male accessory gland secretion is indeed important for achieving the maximal effect on sperm survival. After sperm storage, the queens also provide sperm with secretions from spermathecal glands and we show that these secretions have a comparable positive effect on sperm viability. SDS gels show that the proteomic profiles of accessory gland secretion and spermathecal fluid secretion hardly overlap, which suggests that males and females use different proteins to enhance sperm viability during, respectively, ejaculation and final sperm storage.     

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.     

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

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

Cultivation of epithelia from the secretory coil of the ovine apocrine gland: Evidence of secretory cell function and ductal morphogenesis In vitro

Summary The secretory coil of the ovine apocrine gland is composed predominantly of two cell types, secretory cells lining the lumen and myoepithelial cells adjacent to the basement membrane. The glands synthesize a number of hormones and growth factors, but analysis of the functions of these molecules may be hampered by the mixing of apocrine and sebaceous secretions in the pilary canal. The purpose of this study was to isolate the glands and devise simple culture procedures to facilitate investigations of secretory cell function. The most successful approach involved microdissection of the secretory coils individually from skin biopsies and culture in Dulbecco’s modified Eagle’s medium. After 1–2 wk in medium, cell outgrowths were seen from explants. These consisted predominantly of populations of epithelial cells, many containing granules. Smaller granules were usually concentrated around the cell nuclei and accumulated lipophilic dyes. Large granules were unreactive. Western analysis showed that cells in culture synthesized nerve growth factor-like peptides, a feature consistent with one of the functions of the gland in vivo. When isolated secretory coils were explanted to culture dishes coated with matrigel, highly compact, multilayered masses of cells grew out. Subsequently, tubular structures formed. The observations suggest that some differentiated functions of gland cells were retained in vitro and that the procedures described provide a system for the study of apocrine secretions in isolation from those of other skin glands.     

Spermathecae of Salamandrina terdigitata (Amphibia: Salamandridae): Patterns of sperm storage and degradation

The spermathecae of female Salamandrina terdigitata were observed using light and transmission electron microscopy during the fallspring period of sperm storage and secretory activity and during the summer stasis. When sperm are stored inside the spermathecae, the product synthesized by spermathecal epithelial cells is exported into the lumen, where it bathes the sperm. During sperm storage some spermatozoa undergo degradation by the spermathecal epithelium. This process, which includes sperm capture by the apical microvilli, formation of endocytic vacuoles and production of lysosomes, becomes prominent shortly after oviposition. In many instances, cells filled with vacuolized spermatozoa and/or residual bodies undergo desquamation from the spermathecal epithelium and enter the lumen together with residual sperm. Desquamated cells, together with residual sperm, are a common feature in the spermathecal lumina at the end of the egg-laying season. Concomitant to the activity of the spermathecal epithelium, macrophages move into the spermathecae from the stroma and contribute to the degradation of both the residual sperm and desquamated epithelial cells. As a result of this degradation activity, spermathecae observed during the short summer stasis appear devoid of secretory product and sperm. By late summer, however, the spermathecae already show early signs of an imminent resumption of biosynthetic activity. © 1995 Wiley-Liss, Inc.     

Spermatodesm reorganization in the spermatophore and in the spermatheca of the bushcricket Tylopsis liliifolia (Fabricius) (Orthoptera,Tettigoniidae)

Spermatozoa of Tettigoniidae are usually transferred to the female by means of a spermatophore which is also the site of feather-shaped spermatodesm formation. These spermatodesms are then transferred to a spermatheca, composed of a spermathecal duct and of a seminal receptacle, involved in storing spermatozoa. In order to extend the knowledge about sperm transfer and spermatodesms reorganization in the Tettigoniidae, a morpho-structural investigation was carried out on spermatophore and spermatheca of Tylopsis liliifolia and on the reorganization of the gametes from the spermatophore. Our results show that the spermatodesms undergo disorganization in the spermatophore; unlike other Tettigoniidae, however, feather-shaped spermatodesms are never found. The epithelium of the spermatheca consists of two cell types, the cuticle-forming and the gland cells, with secretory features. The gland cells, absent in the distal tract of the seminal receptacle, release their secretion in a “reservoir” where an efferent duct opens. In the distal tract of the spermathecal duct, adjacent epithelial cells show diversified ultrastructural characteristics whose probable role is discussed. A particular feature of T. liliifolia is the genesis of the feather-shaped spermatodesms in the seminal receptacle. This feature and the peculiar organization of the feather-shaped spermatodesm are a possible autapomorphy of T. liliifolia.     

Development of female accessory glands of Chrysomya putoria (Wiedemann) (Diptera: Calliphoridae) during oogenesis

Females of Chrysomya putoria (Diptera: Calliphoridae) have two sexual accessory glands, which are tubular and more dilated at the distal extremity. The glands open independently into the common oviduct. Two morpho-physiological regions were distinguished in the longitudinal semi-thin sections of the glands. The secretory region is constituted by three layers: a cuticular intima, lining the lumen, followed by a layer of small cells, and then a layer of very large secretory cells. The ductal region of the gland presents only two layers: the cuticular intima and a cellular layer. In both regions a basement membrane is present. Each secretory cell has in its apical region a reservoir, which enlarges throughout oogenesis; in its basal region there is a large nucleus. The ductal cells are cylindrical and smaller than the secretory cells. The glandular secretion is synthesized in the cytoplasm of the secretory cells, stored and/or modified in the reservoir, then drained to the lumen through an end apparatus seen in the apical region of the secretory cell. Histochemical tests indicate that this secretion is a glycoprotein. Measurements of the glands from females at different physiological conditions and fed on different diets correlate with the results obtained for changes in the ovary during oogenesis. Cell number averaged 561.2 ± 77.54 per gland. There was no increase in cell number during oogenesis.     

Sperm storage and degradation in the spermathecae of the salamander Eurycea Cirrigera

Spermathecae are exocrine glands in the roof of the female cloaca that store sperm. Cytological and histochemical data indicate that the one type of secretion into the lumen is a glycoprotein. After a period of stasis in the summer, production of the secretion is initiated in the fall, coincident with an increase in ovarian follicular size. By the time of maximal follicular development and most intense mating activity in March and April, the spermathecal epithelium is filled with secretory granules. The secretory material is released into the lumen, enveloping the sperm. Many sperm remain in the spermathecae after oviposition, and most of these sperm are degraded in the spermathecal epithelium or pass through interruptions in the spermathecal walls caused by desquamation. Sperm in contact with the stromal environment are phagocytized by leukocytes. Some sperm, however, may survive in the lumen until at least the following fall. These sperm retain normal cytology, but whether or not they remain fertile and intact until a subsequent ovipository cycle is unknown.