The organization of the female reproductive system in the parasitic mite Myobia murismusculi (Trombidiformes: Myobiidae)

The general morphology of the internal genital organs in Myobia murismusculi females have been studied by means of light and electron microscopy. It is shown, that their reproductive system is composed of the single ovary, long complex oviduct, cuticular vagina and receptaculum seminis. The apical region of the ovary is formed by the numerous processes of gonadian somatic cells. These cells generate the ovarian sheath and probably take place in oocyte's metabolism. Any nutritive cells in the Myobia ovary are absent. The vitellogenetic oocytes develop in the ovarian pouches like in other trombidiform mites. The oviduct consists of three parts, each of them has a specific ultrastructure and type of secretory product. Two proximal parts of the oviduct produce the egg-shell components, while the third distal part never contains oocytes. It could be suggested that this part produces a special secret onto the oocyte surface to glue the oocytes to the hairs of host.     

Structure of the ovary and mode of oogenesis in a freshwater crab Potamon dehaani

Structure of the adult ovary and oogenetic mode were examined in the freshwater crab Potamon dehaani. An H‐shaped ovary consisting of a pair of long ovarian sacs connected by a narrow bridge tube is located in the cephalothorax on the dorsal side of the stomach. A short oviduct with a seminal receptacle is connected with the posterior end of each ovarian sac, and a genital pore opens on the sternum of the sixth thoracic segment. The ovarian wall consists of a layer of ovarian epithelium that infolds to form a number of oogenetic pouches of various sizes. These are present mainly in the anterior regions of the ovarian sacs, are scarce in the posterior regions of the ovarian sacs, and are absent from the bridge tube. Each oogenetic pouch contains an egg or a relative large oocyte in its lumen. Germaria containing oogonia, very early previtellogenic oocytes, and somatic interstitial cells are located in the ovarian epithelium near the necks of the oogenetic pouches in the anterior regions of the ovarian sacs and are randomly scattered throughout the ovarian epithelium in the posterior regions of the ovarian sacs. In cross section, the germaria appear to be concentrated into a central germarial cluster in the ovarian sac. In the posterior regions of the ovarian sacs, however, the germaria are randomly scattered throughout the ovarian epithelium. An early previtellogenic oocyte leaves its germarium and raises the ovarian epithelium infolds to form a new oogenetic pouch in which it grows to maturity. Mature eggs are ovulated from the oogenetic pouches into the ovarian lumen, transferred from the ovarian lumen into the oviducts, fertilized there by sperm stored in the seminal receptacles, and then oviposited through the genital pores. The female reproductive system is surrounded wholly and tightly by a thin, cellular, membranous sheath, which has often been mistaken as the ovarian epithelium in some decapod crustaceans. J. Morphol. 239:107–114, 1999. © 1999 Wiley‐Liss, Inc.     

Ultrastructural investigations of the female genital system of Epiperipatus biolleyi ( Bouvier 1902) (Onychophora,Peripatidae)

Brockmann, C., Mummert, R., Ruhberg, H. and Storch, V. 1999. Ultrastructural investigations of the female genital system of Epiperipatus biolleyi (Bouvier 1902) (Onychophora, Peripatidae). — Acta Zoologica (Stockholm) 80: 339–349. The female genital system of the neotropical peripatid Epiperipatus biolleyi was examined using transmission and scanning electron microscopy. Special attention is given to the two accessory organs of the paired oviducts: the receptacula seminis and the ovarian funnels (Ovarialtrichter). The latter occur only in the Peripatidae, whereas receptacula seminis may also be present in the Peripatopsidae, the only other family in the Onychophora. The ovarian funnels of E. biolleyi are thin-walled and closed from the haemocoel. This trait has also been reported from Epiperipatus trinidadensis, Macroperipatus torquatus, and Eoperipatus weldoni, whereas other peripatids have ovarian funnels which have been reported to open into the haemocoel. The occurrence of artificially opened ovarian funnels, caused by tissue rupture during specimen preparation, is discussed. The presence of spermatozoa both in the receptaculum seminis and in parts of the uterus of the female examined in this study supports the hypothesis that in E. biolleyi insemination of juvenile females occurs directly via the genital opening. The female contained one unstalked cleavage embryo in each uterus horn. Two features of the uterus were found to be unique to E. biolleyi: (1) a second cell layer overlying the uterine epithelium with a pronounced secretory activity (2) embryos are enclosed in a noncellular coat interspersed with numerous transport vesicles.     

Ultrastructure of receptaculum seminis in Haemaphysalis longicornis (Acari: Ixodidae) in relation to inserted endospermatophore

The receptaculum seminis, opening into the female genital tract, is found only in the metastriate ixodid ticks. An endospermatophore that has been inserted into the female genital aperture at copulation is first stored in the receptaculum seminis, where spermiogenesis is completed before the sperm ascend the oviducts. The receptaculum seminis consists of a simple cuticularized epithelium. Epithelial cells in sexually matured females develop during feeding and become active in secretion. Secretions discharged from epithelial cells are released into the lumen of this organ through the cuticle and may act on the wall of the inserted endospermatophore. The fact that resumption of spermiogenesis (spermateleosis) has already occurred before destruction of the endospermatophore just after copulation suggests that secretions from epithelial cells of the receptaculum seminis are not the trigger of spermateleosis, but a destructive agent of the endospermatophore wall. J Morphol 231:143–147, 1997. © 1997 Wiley-Liss, Inc.     

Effect of blood meal and mating on the genital tract ultrastructure in the female camel tickHyalomma (Hyalomma) dromedarii (Ixodoidea: Ixodidae)

The genital tract ultrastructure in the femaleHyalomma (Hyalomma) dromedarii is described during feeding and mating and up to oviposition. The vagina, consisting of vestibular (VV) and cervical (CV) regions, is formed of an epithelium lined internally with a folded cuticular layer and surrounded externally by muscle layers. These facilitate the passage of endospermatophores containing sperm into the receptaculum seminis (RS), and ova to the exterior. A pair of tubular accessory glands (AG) opening at the junction of VV and CV consist of an epithelial layer of undifferentiated cells. As feeding progresses, these cells synthesise their granular secretion which lubricates the egg surface during its passage through VV. The RS, opening anteriorly into the CV, consists of cuboidal cells lined with a thin cuticular layer. These cells become rich in glycogen and lipid vacuoles, possibly acting as a source of energy for various cell activities including granule synthesis and exocytosis. The granules discharge their contents into intercellular spaces distributed throughout the RS wall and communicate with the main lumen via narrow channels. The cell secretion may dissolve the endospermatophore wall to release sperm, while their lysosome-like structures may function in the breakdown of endospermatophoric material taken up by pinocytosis. The connecting tube (CT) opens into the CV anterodorsally and leads into the common oviduct (COV) posteriorly. It consists of an epithelium lined by a closely-adhering cuticular layer, giving the tube lumen the appearance of an undulate labyrinth with a complicated configuration. No secretory activity in the CT has been observed before, during, or after feeding. The paired, non-cuticular oviducts, extending from the ovary and fusing anteriorly to form COV, consist of an epithelium poor in cell organelles. At the final stages of feeding the cell cytoplasm contains large, phagosomal vacuoles penetrated by sperms, in addition to micropinocytotic vesicles which serve to break down the seminal fluid and other materials. The basal membrane is infolded giving characteristic features, which increase dramatically during oviposition, of epithelia involved in ion and water transport. The oviducal secretion may function as a tanning agent to harden the egg-shell and may also act as a lubricant for egg passage.     

Anatomy and ultrastructure of the female reproductive system of Pleioplana atomata (Platyhelminthes: Polycladida)

The ultrastructure of the female reproductive system of the polyclad flatworm Pleioplana atomata is described. Numerous ovaries are scattered throughout the entire body but are mainly concentrated on the dorsal side. Within an ovary, a germinative zone with oogonia and prefolicular cells is located in the dorsal part of the ovary. The remaining part of the gonad is filled with previtellogenic and early vitellogenic oocytes enwrapped by follicular cells. During previtellogenesis, oocytes produce numerous eggshell globules, which are distributed into the cortical area of the cell in later stages. Eventually, these globules release their contents into the space between the eggshell cover and oolemma. Similar types of globules are also found in others flatworms, and may represent useful phylogenetic characters. Entolecital, vitellogenic oocytes pass to paired uteri, where vitellogenesis is completed. The remainder of the female reproductive system consists of paired thin uterine ducts that join a vagina. The distal part of the long, curved vagina forms a large Lang's vesicle, while the proximal part is connected to a female atrium leading to a female gonopore. We hypothesize that Lang's vesicle functions in the digestion of excess sperm received. Two kinds of different shell (cement) glands that release their secretion into the vagina are identified. Both are unicellular glands and each gland cell connects to the lumen of the vagina via an individual canal. Similar glands in other acotylean polyclads have been implicated in the formation of eggshell covers. J. Morphol. 2009. © 2008 Wiley‐Liss, Inc.     

粉尘螨生殖系统超微结构的透射电镜观察

本研究主要采用透射电镜观察粉尘螨Dermatophagoides farinae (Hughes)生殖系统超微结构。粉尘螨雄性生殖系统是由精巢、 输 精管、 附腺、 射精管、 交配器官及附属交配器官组成。精巢内可同时有精子发育各阶段的细胞。精子无核膜、 核染色质聚集成束、 线 粒体缺乏典型的嵴、 胞质内有平行排列的电子致密薄片等为其特征性结构。雌性生殖系统由交合囊、 交合囊管、 储精囊、 囊导管、 卵 巢、 输卵管、 子宫及产卵管构成。卵巢内可见含多个细胞核的中央细胞, 其周为卵母细胞等生殖细胞。该研究丰富了对粉尘螨生殖系统 结构的认识。     

The fine structure of the ovary of the feather star Nemaster rubiginosa (Echinodermata: Crinoidea)

The outer layer of the crinoid ovary consists of coelomic epithelium, smooth muscles, and nerve cell processes. The middle layer of the ovary contains non-germinal accessory cells, small germinal cells (either oogonia or pre-leptotene primary oocytes), and post-pachytene primary oocytes; all these cells are completely embedded in a haemal matrix of 200 A-diameter granules. The primary oocytes larger than 20mu in diameter have abundant invaginations in the plasma membrane, suggesting uptake of materials from the haemal matrix. The innermost layer of the ovary is a ciliated epithelium lining the cell-free ovarian lumen.     

Histochemical studies on Raillietina (Raillietina) johri (Cestoda: Davaineidae). I. Nonspecific and specific phosphatases.

Certain phosphatases have been localized by histochemical techniques in various tissues of a pigeon cestode, Raillietina (Raillietina) johri. Acid phosphatase (AcPase), alkaline phosphatase (AlPase) and adenosine triphosphatase (ATPase) were present in almost all structures: tegument; subtegumental muscles; subtegumental cells; excretory canal; testes; sperm ductules; vas deferens; cirrus sac; cirrus; ovary; receptaculum seminis; vagina; vitelline gland cells; oocytes; uterus; embryonated eggs. AlPase was absent in parenchyma, spermatocytes, spermatids and spermatozoa. AlPase activity was more intense in the tegument of mature gravid proglottides. AcPase and ATPase were visualized in various stages of spermatogenesis of the parasite. ATPase activity was also observed in chromosomes. 5'-nucleotidase (AMPase) activity was restricted to embryonated eggs only. Functional significance of these phosphatases is discussed.     

Egg-oviduct interaction initiates reproductive behavior

The experiments reported in this paper provide evidence that eggs must pass through the oviducts in order for receptivity to occur after ovulation in the female frog, Rana pipiens. In one experiment, oviductectomized frogs remained unreceptive after ovulation was induced by administration of exogenous pituitary glands, while sham-operates became receptive within 48 hr. Another experiment had four groups of subjects: ovariectomized females, females with oviducts ligated at the ostial end, females with openings in the uteri that prevented eggs from accumulating there, and sham-operated females. Only the last two groups, groups in which eggs could pass through the oviducts, became receptive. In these experiments, receptivity was indicated by absence of the release call during manual clasping of the trunk. Earlier experiments have shown that eggs have to pass through the oviducts in order to become fertilizable. Thus, the passage of eggs through the oviducts provides a mechanism which links the onset of reproductive behavior to the availability of fertilizable gametes.     

Morphology of the female reproductive system of European pea crabs (Crustacea,Decapoda, Brachyura,Pinnotheridae)

Commensal pea crabs inhabiting bivalves have a high reproductive output due to the extension andfecundity of the ovary. We studied the underlying morphology of the female reproductive system in the Pinnotheridae Pinnotheres pisum, Pinnotheres pectunculi and Nepinnotheres pinnotheres using light microscopy and transmission electron microscopy (TEM). Eubrachyura have internal fertilization: the paired vaginas enlarge into storage structures, the spermathecae, which are connected to the ovaries by oviducts. Sperm is stored inside the spermathecae until the oocytes are mature. The oocytes are transported by oviducts into the spermathecae where fertilization takes place. In the investigated pinnotherids, the vagina is of the “concave pattern” (sensu Hartnoll 1968 ): musculature is attached alongside flexible parts of the vagina wall that controls the dimension of its lumen. The genital opening is closed by a muscular mobile operculum. The spermatheca can be divided into two distinct regions by function and morphology. The ventral part includes the connection with vagina and oviduct and is regarded as the zone where fertilization takes place. It is lined with cuticle except where the oviduct enters the spermatheca by the “holocrine transfer tissue.” At ovulation, the oocytes have to pass through this multilayered glandular epithelium performing holocrine secretion. The dorsal part of the spermatheca is considered as the main sperm storage area. It is lined by a highly secretory apocrine glandular epithelium. Thus, two different forms of secretion occur in the spermathecae of pinnotherids. The definite role of secretion in sperm storage and fertilization is not yet resolved, but it is notable that structure and function of spermathecal secretion are more complex in pinnotherids, and probably more efficient, than in other brachyuran crabs. J. Morphol., 2011. © 2010 Wiley‐Liss, Inc.     

Phylogenetic significance of the structure of adult ovary and oogenesis in a primitive chilognathan diplopod,Hyleoglomeris japonica Verhoeff (Glomerida,Diplopoda)

Some histological details of the adult ovary of Hyleoglomeris japonica are described for the first time in the glomerid diplopods. The ovary is a single, long sac-like organ extending from the 4th to the 12th body segment along the median body axis, lying between the alimentary canal and the ventral nerve cord. The ovarian wall consists of a layer of thin ovarian epithelium which surrounds a wide ovarian lumen. A pair of longitudinal “germ zones,” including female germ cells, runs in the lateral ovarian wall. Each germ zone consists of two types of oogenetic areas: 1) 8–12 narrow patch-shaped areas for oogonial proliferation, arranged metamerically in a row along each of the dorsal and ventral peripheries, and 2) the remaining wide area for oocyte growth. Oogonial proliferation areas include oogonia, very early previtellogenic oocytes, and young somatic interstitial cells, among the ovarian epithelial cells. The larger early previtellogenic oocytes in the oogonial proliferation areas are located nearer to the oocyte growth area, and migrate to the oocyte growth area. They are surrounded by a layer of follicle cells and are connected with the ovarian epithelium of the oocyte growth area by a portion of their follicles. They grow into the ovarian lumen, but their follicles are still connected with the oocyte growth area. Various sizes of the previtellogenic and vitellogenic oocytes in the ovarian lumen are connected with the oocyte growth area; the smaller oocytes are connected nearer to the dorsal and ventral oogonial proliferation areas, while the larger ones are connected nearer to the longitudinal middle line of the oocyte growth area. Following the completion of vitellogenesis and egg membrane formation in the largest primary oocytes, the germinal vesicles break down. Ripe oocytes are released from their follicles directly into the ovarian lumen to be transported into the oviducts. Ovarian structure and oogenesis of H. japonica are very similar to those of other chilognathan diplopods. At the same time, however, some characteristic features of the ovary of H. japonica are helpful for understanding the structure and evolution of the diplopod ovaries. Some aspects of the phylogenetic significance in the paired germ zones of H. japonica are discussed. J. Morphol 231:277–285, 1997. © 1997 Wiley-Liss, Inc.     

Oogenesis in Pentastomida

During oogenesis in Pentastomida the oocytes wander from a germinal zone in the mid-dorsal ovarial wall to the lateral pouches of the ovary. The growth phases are initiated during that migration, but not until the lateral pouches are reached does the oocyte grow considerably. A Balbiani body consisting of numerous mitochondria, and various vacuolar inclusions are formed, and at the same time the oocyte bulges out into the hemocoel. Some wall cells form a stalk and cover part of the oocyte towards the hemocoel. During later stages of oocyte growth they withdraw, and the oocyte is separated from the hemocoel only by an extremely thin basement membrane. It seems that the young oocytes obtain nourishment from the ovarial lumen by way of the wall cells. The contents of the lumen in its turn seems to be derived from the hemocoel through the activity of mid-ventral secretory cells. The older oocytes are provided with numerous microvilli, and material for the growth of the oocytes must be taken up directly from the hemocoel. A very dense egg shell is secreted by the oocyte. In nearly mature oocytes the microvilli withdraw from penetrating this shell, and a mucopolysaccharide layer is formed beneath the shell. The mature oocytes in some unknown way make their way to the ovarial lumen whence they are transported to the uterus.     

Allatostatin in ovaries, oviducts, and young embryos in the cockroach Diploptera punctata

The quantity and localization of -Phe-Gly-Leu-amide allatostatins (-F-G-L-amide AST) was determined by ELISA and immunohistochemistry in ovaries and oviducts and in pre-dorsal closure embryos. AST in the cytoplasm of basal oocytes gradually increased from 4 to 35 fmol/ovary pair from the start (day 2) to the completion of vitellogenesis (day 6), then rapidly increased to 121 fmol/ovary pair during choriogenesis. In oviducts, AST-immunoreactivity was found in nerves to the muscle layer and in epithelial cells. AST-immunoreactivity in oviduct epithelial cells increased during vitellogenesis. A marked increase in quantity of AST in oviduct tissue between completion of chorion formation and immediately after ovulation appears to result from AST released from oocytes as they travel down the oviducts because AST content of newly ovulated eggs was 40% lower than late stage chorionated oocytes, and these oocytes released AST when incubated in saline. AST in embryos, localized in yolk cells, decreased as embryos approached dorsal closure. That this material in ovaries and embryos is AST was confirmed by its ability to inhibit JH synthesis in vitro and identification by MALDI-TOF mass spectrometry of a peptide with a mass corresponding to that of a Diploptera punctata AST. These findings indicate likely novel functions for ASTs: facilitation of ovulation and utilization of yolk.     

Morphology of the ovaries of Sphaerodema (Diplonychus) rusticum (Heteroptera,Belostomatidae)

The female reproductive system of Sphaerodema rusticum consists of a pair of ovaries, two lateral oviducts, a median common oviduct, and a median spermatheca. Accessory glands are absent. Each ovary has five free ovarioles branching from the oviduct. Each ovariole consists of a terminal filament, germarium, vitellarium, brown mass, and an exceptionally long pedicel. The terminal filament consists of a central core, interstitial cells, and an outer sheath. In the germarium, which consists of trophic and prefollicular regions, the trophic region or nurse cell chamber is divided into four histologically differentiated zones, distinguished as zones I–IV. Nutritive cords, originating from the posterior end of the trophic core in zone IV extend centrally and join the developing oocytes in the prefollicular chamber and the vitellarium. The compact prefollicular tissue at the base of the trophic core gives rise to prefollicular cells which, after encircling the young oocytes, become modified into follicular epithelial cells, the interfollicular plug, and epithelial plug. The young oocytes descend into the vitellarium and gradually develop into mature oocytes. A compound corpus luteum is observed simultaneously in all the ovarioles of both ovaries after ovulation. Below the epithelial plug there is an accumulation of material, the “brown mass,” which develops cyclically in correlation with the ovulation cycle. Each pedicel stores five mature chorionated eggs ready for oviposition. The epithelium of the anterior region of the pedicel secretes a PAS-positive material. General morphology and histology of the subdivisions of the ovarioles are described.     

Ultrastructure of the eggshell and its formation in Planipapillus mundus (Onychophora: Peripatopsidae)

Although the majority of onychophorans are viviparous or ovoviviparous, oviparity has been described in a number of species found exclusively in Australia and New Zealand. Light microscopy and scanning and transmission electron microscopy were used to examine developing eggs and the reproductive tract of the oviparous Planipapillus mundus. Deposited eggs and fully developed eggs dissected from the terminal end of the uteri have an outer thick, slightly opaque chorion, and an inner thin, transparent vitelline membrane. The chorion comprises an outermost extrachorion, sculptured with domes equally spaced over the surface; a middle exochorion, with pores occurring in a pattern of distribution equivalent to that of the domes of the extrachorion above; and an innermost, thick endochorion consisting of a spongelike reticulum of cavities comparable to the respiratory network found in insect eggs. The vitelline membrane lies beneath the chorion, from which it is separated by a fluid‐filled space. The vitelline membrane tightly invests the developing egg. Examination of oocytes in the ovary and developing eggs at various stages of passage through the uterus indicate that the majority of chorion deposition occurs in the midregion of the uterus, where vast networks of endoplasmic reticulum are present in the columnar epithelium. The vitelline membrane, however, is believed to begin its development as a primary egg membrane, surrounding the developing oocytes in the ovary. The vitelline membrane is transformed after fertilization, presumably by secretions from the anterior region of the uterus; hence, it should be more accurately referred to as a fertilization membrane. Aspects of the reproductive biology of P. mundus are also included. J. Morphol., 2008. © 2008 Wiley‐Liss, Inc.     

Ultrastructure of somatic tissues in the ovaries of a chaetognath (Ferosagitta hispida)

Transmission electron microscopy reveals that the ovaries of Ferosagitta hispida contain four somatic tissues. A myoepithelial ovary wall, continuous with a thin layer of peritoneocytes lining the coelomic cavity, encloses a fluid-filled ovarian space in which oocytes develop. Lamellar extensions of a “follicular reticulum” branch throughout the ovarian space and ensheath developing oocytes. This tissue has been overlooked in most previous studies of chaetognath ovaries. A bipartite oviductal complex extends the length of each ovary just within the lateral ovary wall. It consists of a flattened, blindly ending cellular tube, herein referred to as the cellular sheath, and an enclosed syncytium. Sheath cells secrete an electron-dense product into the ovarian space. Those sheath cells directly bordering the syncytium are contractile and are joined to the to the syncytium by gap junctions and microvillar interdigitations. The syncytium contains a complex of membrane-bounded lumina. The latter sometimes enclose sperm received during mating or ovulated eggs. Thus the syncytium serves both as a seminal receptacle and as a duct for passage of eggs to the outside. Contrary to several classical reports, the cellular sheath and syncytium of the oviductal complex do not separate at ovulation to form a temporary oviductal lumen.     

Fine structure of the female reproductive system in Sobolevicanthus gracilis and Cloacotaenia megalops (Cestoda: Cyclophyllidea)

The fine structure of the female reproductive organs and ducts in two cyclophyllidean cestodes was studied by transmission electron microscopy. All the studied ducts (vitelline, fertilization and vaginal ducts, and oviducts) as well as the uterine and ootype walls have the luminal surface elevated to form apical cytoplasmic lamellae and are surrounded by bands of circular muscules, which are attached to the basement layer. The structure of the studied ducts epithelium corresponds to the pattern described in other ducts of the cestode genital system, namely a nucleate syncytial layer. An exception is the vagina Sobolevicanthus gracilis, in which the surface is lined with not numerous atypical microtriches. The comparison of our results and the data reported for other species is given. It is found out that in different species of cestodes forming oligolecital eggs, there are observed various uterine structures and diverse contacts between the eggs capsules and the uterine epithelium. The formation of special structures and contacts is the evolutionary way from the extrauterine type to more progressive intrauterine type of embryo development in cestodes.     

Embryogenesis and the first-stage larva of Thelazia lacrymalis

The female reproductive system of Thelazia lacrymalis (Nematoda: Thelaziidae) was investigated by light and scanning electron microscopy (SEM) with regard to the developmental stages and the stage deposited by the gravid nematode. Female T. lacrymalis have a didelphic and opisthodelphic type of reproductive system with paired ovaries, oviducts and uteri and a single vagina and vulva. Round and spindle-shaped primary oocytes are documented within the ovaries and oviducts, respectively. The distal part of each uterus provides a fertilization chamber filled with spermatozoa, followed by a sphincter-like part. Further anterior, the uteri broaden gradually containing dividing zygotes, small and large morulae, tadpole-stage embryos and horseshoe-shaped embryos which increase in length and become slimmer forming pretzel-stage embryos and larvae rolled up. The larvae stretch gradually and finally lie straight but still covered with their egg membrane in the vagina. The egg membrane encloses the whole larva and is enlarged at the pointed tail of the larva forming a bulb. At the SEM level, the first-stage larva is shown to have a terminal mouth and three hooks directed posteriorly and a striated cuticle. As morphologically identical larvae were also found in lavages of the conjunctival sac of horses infected with T. lacrymalis, this nematode species can be described as ovoviviparous.