Anatomy and ultrastructure of the reproductive systems ofAcarus siro (Acari: Acaridae)

Anatomy and ultrastructure of the female and male reproductive system inAcarus siro L. were investigated by light and electron microscopy. The female system consists of paired ovaries of nutrimentary type in which oogonia and oocytes are connected by bridges with a large central cell. The oviducts empty into the uterus, which passes into preoviporal duct lined bycuticle, and opening as a longitudinal slit (oviporus). An elongated accessory gland composed of one type of secretory cell is located along each oviduct. The copulatory opening occurs at the posterior margin of the body and leads, via the inseminatory canal, to the receptaculum seminis, consisting of the basal and saccular part. Both inseminatory canal and basal part of receptaculum seminis are lined by cuticle, whereas the wall of the sac is formed by cells covered only by long, numerous microvilli. The basal part of the receptaculum seminis joins the ovaries via two lumenless transitory cones.The male reproductive system contains paired testes, in which spermatogonia tightly surround the central cell. The proximal part of the paired vasa deferentia serves as a sperm reservoir, while the distal one has a glandular character. An unpaired, cuticle-lined ejaculatory duct opens into the apex of the aedeagus. The single accessory gland is located asymmetrically at the level of, or slightly posterior to, coxae IV.The structure of the genital papillae, which are topographically related to the genital opening in both sexes, is also briefly described.     

粉尘螨生殖系统形态学研究

粉尘螨Dermatophagoides farinae是一种重要的医学螨类。本文用光镜和扫描电镜研究了粉尘螨雌雄生殖系统的形态和结构。结果表明：雄性生殖系统由睾丸、输精管、 附腺、射精管、阳茎及附属交配器官组成。睾丸位于血腔末端,不成对,精原细胞、精母细胞和精子依照精子发育的顺序有规则地分布在其内部。雌性生殖系统包括交配孔、囊导管、储精囊、囊导管、卵巢、输卵管、子宫、产卵管及产卵孔,其中卵巢由一个中央营养细胞和围绕其周围的卵母细胞组成。     

Evaluating the post‐copulatory sexual selection hypothesis for genital evolution reveals evidence for pleiotropic harm exerted by the male genital spines of Drosophila ananassae

The contemporary explanation for the rapid evolutionary diversification of animal genitalia is that such traits evolve by post‐copulatory sexual selection. Here, we test the hypothesis that the male genital spines of Drosophila ananassae play an adaptive role in post‐copulatory sexual selection. Whereas previous work on two Drosophila species shows that these spines function in precopulatory sexual selection to initiate genital coupling and promote male competitive copulation success, further research is needed to evaluate the potential for Drosophila genital spines to have a post‐copulatory function. Using a precision micron‐scale laser surgery technique, we test the effect of spine length reduction on copulation duration, male competitive fertilization success, female fecundity and female remating behaviour. We find no evidence that male genital spines in this species have a post‐copulatory adaptive function. Instead, females mated to males with surgically reduced/blunted genital spines exhibited comparatively greater short‐term fecundity relative to those mated by control males, indicating that the natural (i.e. unaltered) form of the trait may be harmful to females. In the absence of an effect of genital spine reduction on measured components of post‐copulatory fitness, the harm seems to be a pleiotropic side effect rather than adaptive. Results are discussed in the context of sexual conflict mediating the evolution of male genital spines in this species and likely other Drosophila.     

Four new species of the genus Perepsilonema Lorenzen, 1973 (Nematoda,Epsilonematidae) from the bay of Calvi (Corsica,Mediterranean)

Four new species of the genus Perepsilonema are described. They are characterized by general habitus, cuticular ornamentation, shape and sexual dimorphism of the amphideal fovea, shape and number of ventral copulatory spines and the shape of the spicules. The female genital system comprises two reflexed ovaries, a left and a right spermatheca and a median uterine chamber filled with sperm cells. The distal part of the uteri have secretory wall cells which contain in some specimen spherical refractive bodies.     

How the length of genital parts affects copulation performance in a carabid beetle: implications for correlated genital evolution between the sexes

To identify factors leading to the correlated evolution of exaggerated male and female genitalia, we studied the effects of the variable dimensions of corresponding functional genital parts (male copulatory piece and female vaginal appendix) on copulatory performance in the polygamous carabid beetle Carabus (Ohomopterus) maiyasanus. We used mating pairs of individuals from two populations to increase the variances in genital dimensions and determined the copulation performance (insemination and spermatophore replacement, and copulation time) in single‐ and double‐mating situations. In single mating, insemination success was not affected by genital dimensions, although the copulation time was significantly shorter when the male aedeagus was longer. In the double‐mating experiment, insemination and replacement of spermatophores by the second male succeeded more frequently when the copulatory piece was shorter and the vaginal appendix was longer, and when the difference between the length of the copulatory piece and the vaginal appendix was smaller. Thus, a matching of the corresponding genital parts between the sexes increases the male's reproductive success in sperm competition, but elongation of the copulatory piece cannot be explained simply by the improvement in male reproductive success. We discuss possible factors for the elongation of genital parts in terms of sexual conflict and reproductive interference through interspecific copulation.     

Structure of the testis and genital duct of freshwater stingray, Himantura signifer (Elasmobranchii: Myliobatiformes: Dasyatidae)

The light microscopic structure of the testis and genital duct system of the freshwater stingray Himantura signifer was observed. The testis is composed of lobes having numerous spermatocysts in a dorsoventral zonated arrangement. The germinal papilla at the middorsal surface of the testicular lobe is the origin site of spermatocyst development, where mesenchymal-like cells are predominantly found. The association of a Sertoli cell precursor with a spermatogonium marks the onset of spermatocyst formation and development. The newly formed spermatocysts at the dorsal end of the germinal zone replace the older ones, which are sequentially moved to the ventral side and are termed spermatogonial, spermatocyte, spermatid, spermatozoal, and degenerate zones. In the degenerate zone, the spermatocysts deteriorate after releasing the spermatozoa into the intratesticular duct, where they are further transported through the extratesticular duct system and finally stored at the seminal vesicle. The epithelial lining of the genital duct is a pseudostratified ciliated columnar with no muscular layer underneath; thus, sperm are conveyed through ciliary activity. The interesting features of the present study are the finding of mesenchymal-like cells in the germinal papilla and the nonaggregated formation of sperm in the seminal vesicle.     

Ultrastructure of the male genital tract,spermatogenesis and spermatozoa of hattena cometis domrow (Acari: Gamasida: Ameroseiidae)

The ameroseiid mite Hattena cometis has a male genital system that consists of an unpaired, u‐shaped testis and paired deferent ducts leading into an unpaired accessory genital gland and ejaculatory duct. The genital opening is located anteriorly immediately in front of the sternal shield. Spermatogenesis is simple, probably due to the haploid nature of the male. Eight stages of spermatogenesis could be roughly distinguished. Mature spermatozoa as found in the deferent duct lumen are peculiar in having a bisected nucleus and numerous peripheral flat chambers, which were formed from indentations of the plasmalemma. In inseminated females, spermatozoa were observed in the syncytial tissue of the sperm access system and in the somatic cells of the ovary. These spermatozoa have achieved a new structure, i.e., an electron‐dense plate dividing the cell into two unequal halves. The dense plate has an intricate substructure. Its function is unknown. These sperm cells are considered to represent capacitated spermatozoa. The peripheral chambers are reduced in number inside the female. Similar sperm cells, containing a dense plate, were seen in vacuoles within the epithelium of the deferent duct of one male. These cells are evidently under destruction, but before being completely dissolved had undergone a development leading beyond that of the mature sperm cells found in the deferent duct. Apparently, entering the cell of the deferent duct epithelium or the syncytium tissue triggers the production of the dense plate (or the capacitation process). Our observations are compared with results obtained from other anactinotrichid Acari, mainly Gamasida, and confirm and complete the interpretation of the correlated evolution of components of gamasid reproductive systems. J. Morphol. 274:1010–1025, 2013. © 2013 Wiley Periodicals, Inc.     

Complex genital system of a haplogyne spider (Arachnida, Araneae, Tetrablemmidae) indicates internal fertilization and full female control over transferred sperm

The female genital organs of the tetrablemmid Indicoblemma lannaianum are astonishingly complex. The copulatory orifice lies anterior to the opening of the uterus externus and leads into a narrow insertion duct that ends in a genital cavity. The genital cavity continues laterally in paired tube-like copulatory ducts, which lead into paired, large, sac-like receptacula. Each receptaculum has a sclerotized pore plate with associated gland cells. Paired small fertilization ducts originate in the receptacula and take their curved course inside the copulatory ducts. The fertilization ducts end in slit-like openings in the sclerotized posterior walls of the copulatory ducts. Huge masses of secretions forming large balls are detectable in the female receptacula. An important function of these secretory balls seems to be the encapsulation of spermatozoa in discrete packages in order to avoid the mixing of sperm from different males. In this way, sperm competition may be completely prevented or at least severely limited. Females seem to have full control over transferred sperm and be able to express preference for spermatozoa of certain males. The lumen of the sperm containing secretory balls is connected with the fertilization duct. Activated spermatozoa are only found in the uterus internus of females, which is an indication of internal fertilization. The sperm cells in the uterus internus are characterized by an extensive cytoplasm and an elongated, cone-shaped nucleus. The male genital system of I. lannaianum consists of thick testes and thin convoluted vasa deferentia that open into the wide ductus ejaculatorius. The voluminous globular palpal bulb is filled with seminal fluid consisting of a globular secretion in which only a few spermatozoa are embedded. The spermatozoa are encapsulated by a sheath produced in the genital system. The secretions in females may at least partly consist of male secretions that could be involved in the building of the secretory balls or play a role in sperm activation. The male secretions could also afford nutriments to the spermatozoa.     

Fine structure of the genital system in the bee parasite,Varroa jacobsoni (Gamasida: Dermanyssina) with remarks on spermiogenesis,spermatozoa and capacitation

The female genital tract ofVarroa jacobsoni is composed of a sperm-access system comprising paired solenostomes located between coxa III and IV, paired tubules, paired rami, an unpaired sperm duct, and an unpaired spermatheca. Another part of the female genital system is confined to egg development and oviposition. It is composed of an ovary (s.str.), in which oocytes mature, and a lyrate organ functioning as a nutrimentary structure. Both compartments, regarded as parts of the gonad, are connected by a region named the camera spermatis. This part is also in continuity with the oviduct I, which is provided with a muscular layer and numerous nerve endings. The following cuticle-lined oviduct II leads to the genital orifice through which the eggs are deposited. The fine structure of all these parts is described. Attention is drawn especially to the peculiar spermatheca which contains the inner cells which are thought to connect by way of free cells with a specialized region of the camera spermatis, thus establishing a cellular bridge through which penetration of capacitated spermatozoa into the ovary s.str. may occur. Lyrate organ and oocytes are connected via intercellular bridges/nutritive cords and are thus comparable to the telotrophic ovarioles of certain insects.The male genital system, composed of unpaired testis, paired vasa deferentia, unpaired accessory gland and ductus ejaculatorius, is described ultrastructurally. Spermiogenesis occurs in cysts and spermatozoa belong to the ribbon type. The vasa deferentia are provided with a muscular layer. For the first time receptors are detected in the proximal part or the ductus ejaculatorius. The accessory gland produces a proteinaceous secretion.Spermatozoa were observed in the female rami and spermatheca. Only in the latter were elongated, capacitated spermatozoa seen.     

Increase of cortisol levels after temperature stress activates dmrt1a causing female‐to‐male sex reversal and reduced germ cell number in medaka

In vertebrates, there is accumulating evidence that environmental factors as triggers for sex determination and genetic sex determination are not two opposing alternatives but that a continuum of mechanisms bridge those extremes. One prominent example is the model fish species Oryzias latipes which has a stable XX/XY genetic sex determination system, but still responds to environmental cues, where high temperatures lead to female‐to‐male sex reversal. However, the mechanisms behind are still unknown. We show that high temperatures increase primordial germ cells (PGC) numbers before they reach the genital ridge, which, in turn, regulates the germ cell proliferation. Complete ablation of PGCs led to XX males with germ cell less testis, whereas experimentally increased PGC numbers did not reverse XY genotypes to female. For the underlying molecular mechanism, we provide support for the explanation that activation of the dmrt1a gene by cortisol during early development of XX embryos enables this autosomal gene to take over the role of the male determining Y‐chromosomal dmrt1bY.     

Female behaviour and the interaction of male and female genital traits mediate sperm transfer during mating

Natural selection and post‐copulatory sexual selection, including sexual conflict, contribute to genital diversification. Fundamental first steps in understanding how these processes shape the evolution of specific genital traits are to determine their function experimentally and to understand the interactions between female and male genitalia during copulation. Our experimental manipulations of male and female genitalia in red‐sided garter snakes (Thamnophis sirtalis parietalis) reveal that copulation duration and copulatory plug deposition, as well as total and oviductal/vaginal sperm counts, are influenced by the interaction between male and female genital traits and female behaviour during copulation. By mating females with anesthetized cloacae to males with spine‐ablated hemipenes using a fully factorial design, we identified significant female–male copulatory trait interactions and found that females prevent sperm from entering their oviducts by contracting their vaginal pouch. Furthermore, these muscular contractions limit copulatory plug size, whereas the basal spine of the male hemipene aids in sperm and plug transfer. Our results are consistent with a role of sexual conflict in mating interactions and highlight the evolutionary importance of female resistance to reproductive outcomes.     

Expression of a cell surface protein during morphogenesis of the reproductive system in Manduca sexta embryos

New antibody markers have allowed more refined examinations of embryogenesis. Features are being found that were overlooked in whole and sectioned embryos stained with traditional histochemical labels. Two monoclonal antibodies that recognize two different cell surface proteins in Manduca sexta label cells of the developing reproductive system. These specific immunolabels reveal that during a brief period of Manduca embryogenesis, rudiments of both male and female genital ducts are present in a single embryo. This transient phase of genital differentiation parallels the transient indifferent stage known to occur during development of reproductive systems in many vertebrate embryos. At the end of this indifferent stage, one of the two pairs of genital ducts retracts and degenerates. The dynamic expression of the two surface proteins on cells involved in morphogenesis of both the female and male reproductive systems also suggests that these proteins are important in orchestrating the specific cellular interactions that occur between mesodermal cells of the genital ducts and the nearby ventral ectoderm.     

Cell lineage restrictions in the genital disc ofDrosophila revealed byMinute gynandromorphs

Summary The genital imaginal disc ofDrosophila differentiates the terminalia, i.e. the genitalia and analia, of both sexes. It represents a composite anlage, containing a female genital primordium, a male genital primordium and an anal primordium. In normal males and females, only one of the two genital primordia differentiates; the other is developmentally repressed. Therefore, cell-lineage relationships between the male and female genital primordia can only be studied in sexual mosaics which differentiate female and male cells. We producedMinute (M)non-Minute(M⁺) gynandromorphs and selected those with sexually mosaic terminalia for a cell-lineage analysis. In these mosaics, either the male (XO) or female (XX) cells wereM ⁺ and thus had a growth advantage. The differential growth rates served as a tool to detect clonal restrictions. In control gynandromorphs (M ⁺M ⁺), the amount of female genitalia differentiated was largely independent of the amount of male genitalia present. In contrast, male and female anal structures, as a rule, added up to one full set. The same was true for the experimentalMM ⁺ gynandromorphs, but the contribution ofXX andXO cells to mosaic terminalia changed drastically due toM ⁺ cells competing successfully against the more slowly growingM cells. Specific subsamples ofMM ⁺ gynandromorphs showed thatM cells in a non-mosaic primordium are shielded from cell competition taking place in the neighbouring mosaic primordium. We conclude that the three primordia of the genital disc represent developmental compartments. In the genital primordia, even developmentally repressedM ⁺ cells compete successfully against developmentally activeM cells.     

Copulatory Plug Displacement Evidences Sperm Competition in Lemur catta

Male displacement of copulatory (sperm) plugs from female vaginas provides further evidence for sperm competition in ring-tailed lemurs (Lemur catta), a gregarious prosimian species with a multimale, multifemale mating system. During two mating seasons, I studied two groups of free-ranging ring-tailed lemurs on St. Catherines Island, GA, USA. I observed 22 mating pairs in which males achieved penile intromission. Copulatory plug displacement by males occurred in 9 cases. Plugs were displaced during copulation by male penes upon withdrawl following deep vaginal thrusting. In every case of copulatory plug displacement, the male displacing a plug mated to ejaculation with the estrous female. In a mating system in which females typically mate with more than one male during estrous, often in succession, copulatory plug displacement may function to disrupt or preclude other males' successful insemination of estrous females. The effects of sperm plug displacement on paternity in Lemur catta are unknown, as no study had heretofore documented copulatory plug displacement in this species. The first-male mating advantage suggested for Lemur catta should be re-evaluated where mating order is known, and copulatory plug displacement during mating, or lack thereof, is identified. Because there is a tendency for first-mating males to mate-guard for longer periods of time in Lemur catta, the latency period between the first mate's ejaculation and that of subsequent mates may be an important determinant of male fertilization success.     

Male genital organs in the eulittoral meiofaunal polychaete <Emphasis Type="Italic">Stygocapitella subterranea</Emphasis> (Annelida,Parergodrilidae): ultrastructure,functional and phylogenetic significance

The marine interstitial polychaete Stygocapitella subterranea is characterized by aberrant morphological and biological traits resembling those of clitellates and Hrabeiella periglandulata, a terrestrial polychaete species. Although clearly related to the terrestrial Parergodrilus heideri, there are distinct differences in their morphology. An ultrastructural study of the male genital organs was undertaken to look for common apomorphic features in Parergodrilidae, to find structural evidence for clarifying their reproductive biology and mode of sperm transfer. Finally it should be elucidated whether a supposed sister-group relationship of Parergodrilidae and Orbiniidae based on molecular evidence can be supported by morphological characters as well. In S. subterranea the male organs consist of an unpaired seminal vesicle, a pair of sperm ducts and two large tube-like prostate glands. These glands constitute the distal parts of the gonoducts and open ventrally on a small genital papilla in chaetiger 9. True copulatory organs or organs for storage of mature sperm are lacking. The seminal vesicle is a coelomic cavity composed of two apposed coelomic linings supplied with blood spaces. The testes are found ventrally. The prostate glands are covered by a single layer of muscle fibres running in a longitudinal/spiral direction along the gland. There are no signs of spermatophore formation in any part of the male system. Since females always carry sperm, pseudocopulation can be excluded and the likelihood of either direct transfer of sperm or hypodermic injection is discussed. The structure of genital organs reveals similarities to those of P. heideri. Gonochorism, paired seminal vesicles and two pairs of male gonoducts opening in chaetigers 9 and 10 with a distal glandular part most likely belong to the ground pattern of Parergodrilidae. The observations confirm that consistencies with either clitellates or H. periglandulata are the result of convergent evolutionary events. On the other hand, the relationship of Parergodrilidae to an orbiniid/questid clade receives support from the present data. This paper is dedicated to our scientific teacher, Professor Wilfried Westheide, who made significant contributions to the reproductive biology of interstitial polychaetes, on the occasion of his 70th birthday.     

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。     

Ultrastructure of the reproductive system of the house dust mitesDermatophagoides farinae andD. pteronyssinus (Acari,Pyroglyphidae) with remarks on spermatogenesis and oogenesis

Several parts of the reproductive system of both sexes ofDermatophagoides farinae andD. pteronyssinus are investigated and compared by light-, scanning electron-, and transmission electron microscopy. New techniques have been employed for scanning of the internal structures of these mites. The male reproductive system consists of unpaired testis, paired vasa deferentia, an accessory gland, ejaculatory duct, and copulatory organ. The female reproductive system consists of bursa copulatrix, ductus bursae, receptaculum seminis, paired ducti receptaculi, ovaries, oviducts, one chorion gland, ovipositor, and oviporus. Testis as well as ovaries are characterized by syncytia of nutritional function. The specifics of spermatogenesis are discussed in connection with sperm transfer. Similarities between the construction of the ovaries and oogenesis in astigmatic mites and telotrophic meroistic ovaries in insects are discussed.     

Copulatory mechanism in a sexually cannibalistic spider with genital mutilation (Araneae: Araneidae: Argiope bruennichi)

Genitalia are among the fastest evolving morphological traits as evidenced by their common function as diagnostic traits in species identification. Even though the main function of genitalia is the successful transfer of spermatozoa, the presence of diverse structures that are obviously not necessary for this suggests that genitalia are a target of sexual selection. The male genitalia of many spider species are extremely complex and equipped with numerous sclerites, plates and spines whose functions are largely unknown. Selection on male genitalia may be particularly strong in sexually cannibalistic spiders, where mating success of males is restricted to a single female. We investigated the copulatory mechanism of the sexually cannibalistic orb weaving spider Argiope bruennichi by shock freezing mating pairs and revealed a complicated interaction between the appendices and sclerites that make up the male gonopods (paired pedipalps). The plate that covers the female genital opening (scape) is secured between two appendices of the male genital bulb, while three sclerites that bear the sperm duct are unfolded and extended into the female copulatory opening. During copulation, females attack and cannibalise the male and males mutilate their genitalia in about 80% of cases. Our study demonstrates that (i) genital coupling is largely accomplished on the external part of the female genitalia, (ii) that the mechanism requires an interaction between several non-sperm-transferring structures and (iii) that there are two predetermined breaking points in the male genitalia. Further comparative work on the genus Argiope will test if the copulatory mechanism with genital mutilation indeed is an adaptation to sexual cannibalism or if cannibalism is a female counter adaptation to male monopolisation through genital plugging.     

A new interpretation of the female genitalia in Macrocyclops albidus (Copepoda, Cyclopidae)

The female genital structures of Macrocyclops albidus (Cyclopidae, Eucyclopinae) were studied using light and electron microscopy. The results confirm that the exterior genital area shows only a copulatory pore, located anteromedially on the ventral face of the genital double-somite, and paired gonopores (not directly visible), situated laterally under the P6 plates. An internal seminal receptacle, composed of several parts, is connected to the gonopores by ventro-lateral cuticular extensions or seminal ducts. The lateral site of communication shows a complex set of connections between the seminal receptacle and the oviducts (via the egg-laying ducts). The structure until now designated as ‘transverse ducts’, visible by transparency on the ventral face, is in fact constituted of internal cuticular thickenings resulting of the fusion of the 6th thoracic somite and the 1st abdominal somite forming the genital double-somite and appearing externally as a part of the suture line; the term ‘suture cord’ is proposed to designate it. The functioning of the system is explained.