Functional morphology of the sperm-containing chambers of the sea slug Okenia polycerelloides in the context of sexual selection

Sea slugs are interesting models to study post-copulatory sexual selection in simultaneous hermaphrodites due to the enormous variation of their reproductive systems. However, the knowledge of the functional morphology of their reproductive system is limited to few species, and it is rarely discussed in the context of sexual selection theory. In this study, we investigated the functional morphology of the sperm-containing chambers (i.e., ampulla, seminal receptacle, and bursa copulatrix) of the reproductive system of Okenia polycerelloides (Ortea & Bouchet, 1983), based on light, confocal, and electron microscopy. Although the morphology of the ampulla is similar to other species, indicating that it is a site for autosperm storage, we found some sperm facing the ampullar epithelium, a feature commonly regarded as characteristic of the seminal receptacle of sea slugs. The seminal receptacle of O. polycerelloides showed secretory activity and contained sperm with distribution and orientation suggestive of stratification of allosperm from distinct mating events, a feature that would affect sperm competition. The bursa copulatrix had epithelial cells with secretory and absorptive characteristics, and contained degraded sperm and yolk granules within its lumen. Comparative analyses of the contents of each organ demonstrated that sperm digestion occurs in the bursa copulatrix and affects sperm heads first, changing their morphology from slender and curved to shorter and ellipsoid before complete lysis. Although digestion and absorption of surplus sperm are currently the main hypothesized functions for the bursa copulatrix, its role in cryptic female choice should not be ruled out. The close structural connection between the seminal receptacle and bursa copulatrix, as well as their muscular walls, would enable control over the fate of the sperm received in each mating event, that is, storage or digestion.     

斑衣蜡蝉雌性生殖系统超微结构

【目的】明确斑衣蜡蝉Lycorma delicatula雌成虫生殖系统整体形态及超微结构特征,为蜡蝉总科昆虫分类及系统发育探讨提供更多形态学证据。【方法】采用光学显微镜与透射电子显微镜,观察斑衣蜡蝉雌成虫生殖系统整体形态和各主要器官的超微结构。【结果】斑衣蜡蝉雌成虫生殖系统主要包括1对卵巢、1个中输卵管、1个交配囊、1个交配囊管、1个前阴道、1个后阴道、1个受精囊、1个受精管和2根受精囊附腺。卵巢为端滋式,由14根卵巢小管组成,卵室由固有膜、滤泡细胞和卵细胞组成,卵巢小管中的滋养细胞清晰可见;中输卵管位于前阴道基部,由中输卵管腔、上皮细胞、肌肉鞘和基膜组成;交配囊膨大呈圆球状,囊壁由上皮细胞、肌肉层和基膜组成;交配囊管呈圆柱状,连接交配囊和后阴道,由肌肉鞘、上皮细胞层和管腔组成;前、后阴道超微结构相似,主要由肌肉鞘、基膜、上皮细胞和管腔组成,但后阴道上皮细胞细胞核周围存在分泌颗粒,且管腔内有大量微绒毛,而前阴道壁内包含有大量囊泡结构;受精管从中输卵管末端延伸至受精囊,由基膜、厚层肌肉鞘和管腔组成;受精囊为受精管近末端略膨大的囊状结构,由肌肉鞘、基膜、上皮细胞和囊腔构成;雌性受精囊附腺着生于受精囊末端,为均匀的螺旋管状,主要由肌肉层、上皮细胞层和附腺中心管腔组成。【结论】斑衣蜡蝉雌性生殖系统与已报道的蜡蝉总科其他类群的雌性生殖系统结构相似,但卵巢小管数目有差异;蝉亚目中不同总科雌成虫雌性附腺与受精囊附腺的形态特征存在明显区别;斑衣蜡蝉雌性生殖系统超微结构与叶蝉总科和沫蝉总科昆虫也存在部分差异。这些差异是否可以作为头喙亚目高级阶元的划分依据仍有待于进一步研究。     

The morphology of the phallosome and accessory gland material transfer during copulation in the blowfly,Lucilia cuprina (Insecta,Diptera)

Summary The mechanism of insemination in Lucilia cuprina is described. There are four duct openings on the phallosome. The terminal pore is continuous with the internal reproductive organs. The sub-terminal pore and paired, lateral pores are the openings of ducts formed by cuticular invaginations. The male accessory gland material appears to be re-routed from the terminal pore into the ducts of the lateral barbs once sperm transfer is complete. A novel means of deposition of the male accessory material is described, in which paired barbs on the phallosome tear the cuticle of thick-walled pads in the bursa copulatrix. After mating, accessory material is found within intra-cuticular spaces in the bursa copulatrix, which appear to become sealed at the points of penetration by plugs or scars.     

Reproductive morphology,copulation, and inter-populational variation in the diamondback moth,Plutella xylostella (L.) (Lepidoptera: Plutellidae)

Copulatory mechanisms and internal reproductive systems of male and female Plutella xylostella (L.) were investigated. Both male and female specimens exhibited characteristics typical of ditrysian Lepidoptera, with some peculiarities. Female structures appear to be relatively simple: the bursa copulatrix lacks cuticular signa within, the spermatheca lacks lagenar arms exhibited in some Lepidoptera, and colleterial glands have secretory system and reservoir combined. Male accessory gland ducts are joined distally, a condition that has not been described in other Lepidoptera. Genitalia of both sexes appear simple; females possess a posterior cuticular extension of abdominal sternites that houses the bursal duct and accomodates the needle-like aedeagus of a male during copulation. Despite the apparent simplicity, configuration of genitalia is highly specific in shape, angle, and size, which is likely to be important in maintaining reproductive isolation within the species. However, a population of P. xylostella from Australia showed some modification to both aedeagal and bursal structures. These variations suggest some important implications for considering species with worldwide distributions.     

Angiotensin I-converting enzyme (ACE) activity of the tomato moth, Lacanobia oleracea: changes in levels of activity during development and after copulation suggest roles during metamorphosis and reproduction

Angiotensin I-converting enzyme (ACE) is a dipeptidyl carboxypeptidase that removes C-terminal dipeptides from relatively short oligopeptides, usually smaller than 15 amino acids. In mammals, the enzyme has several important roles in the metabolism of vasoactive peptides, but its physiological role in insects is not fully understood. We now report the properties of an ACE in a lepidopteran species (the tomato moth, Lacanobia oleracea) and suggest new physiological roles for the enzyme in this insect. ACE activity increases four-fold during the last stadium and in early pupae, a rise which, in its timing, is similar to what has been observed previously in the transition of larva to pupa in Drosophila melanogaster. This suggests that the increase in ACE activity might be of general importance for peptide metabolism during metamorphosis in holometabolous insects. High levels of ACE activity were found in the haemolymph of sixth stadium larvae and adult insects, and in the reproductive tissues of both male and female adults. Almost all of the ACE activity in the reproductive tissues was found in the accessory glands of the male and the spermatheca and bursa copulatrix of the female. The decline in accessory gland ACE in mated males and the concomitant rise in ACE activity in the spermatheca and bursa copulatrix of the female suggested the transfer of ACE from the male to the female during copulation. Using several convenient peptides as substrates, we have shown that the spermatophore/bursa copulatrix taken from mated female insects possess an aminopeptidase, a carboxypeptidase and a dipeptidase, in addition to high levels of ACE. These peptidases might be involved in the breakdown of proteins to peptides and eventually to amino acids in the spermatophore. Evidence for such a proteolytic pathway and its role in providing substrates for the TCA cycle has been obtained previously in a study of reproduction in Bombyx mori.     

Reproductive morphology, mating behavior, and spawning ecology of cephalaspid sea slugs (Aglajidae and Gastropteridae)

Abstract. Several recent experimental studies have assessed the behavioral reproductive ecology of four opisthobranch species belonging to the families Aglajidae and Gastropteridae (Cephalaspidea). In order to extend the scope of this earlier work, here we document the reproductive morphology and behavior of 36 taxa belonging to the same two sea slug families. Measurements focus on insemination patterns, spawning characteristics, as well as the size and shape of male and female reproductive organs (i.e., prostate, penis, bursa copulatrix, seminal receptacle). We found that behavioral and morphological traits vary considerably between species, even between closely related taxa. Mating behavior covers the full spectrum from unilateral to simultaneously reciprocal insemination and includes putatively antagonistic mechanisms such as hypodermic injection in some cases. With respect to morphology, particularly great variation was present in traits related to the male sexual function, such as prostate size and penis type. In traits associated with the female function, noticeable variation in those traits accessible for measurement was limited to the relative size of the sperm‐digesting bursa copulatrix. These data provide the basis for future comparative analyses on the evolution of reproductive characters in a phylogenetic context. We further explore the relationship between characteristics and body size. Our data corroborate previous findings that in species with exclusively planktotrophic development, investment per embryo (using mean egg diameter per species as a proxy) is independent of mean species body size. In contrast to egg size, spawn weight and the number of eggs per spawn tightly increased with body size, both within and among species. Hence, larger individuals among planktotrophs increased reproductive investment by increasing offspring quantity rather than investing more into each single offspring. Comparably large eggs with few eggs per spawn in our only two species with facultatively intracapsular metamorphosis are also consistent with findings in other opisthobranchs.     

Arnhemtrachia ramingining: a new genus and species of land snail from Arnhem Land,Australia (Stylommatophora: Camaenidae)

The new, monotypic camaenid genus Arnhemtrachia is described for the new species Arnhemtrachia ramingining from Ramingining, Arnhem Land, Northern Territory, Australia with additional records from the Gove Peninsula in eastern Arnhem Land. This camaenid is characterised by a combination of distinct morphological features, such as a small, discoidal, keeled shell with regular axial ribs, open umbilicus and simple, rounded aperture as well as a reproductive system with a long epiphallus possessing a long flagellum and absence of a penial sheath, with a flap on the penial wall which transverses longitudinal pilasters, and a long, simple bursa copulatrix. Based on comparative morphology, Arnhemtrachia reveals affinities with Trozena morata Iredale, 1838 from NE Queensland.     

Functional morphology of the gonoduct of the viviparous teleost Poeciliopsis gracilis (Heckel, 1848) (Poeciliidae)

Female teleosts do not develop Müllerian ducts; consequently, the ovary of teleosts contains two zones: germinal and gonoduct. The gonoduct lacks germinal cells, but has relevant functions in the reproductive process. We describe the functional morphology of the gonoduct in the viviparous teleost Poeciliopsis gracilis during nongestation and gestation stages. This study tests the hypothesis that the gonoduct functions as a barrier between the germinal zone and the exterior. By providing information about morphology and function of the gonoduct we show that this part of the ovary has an essential role in the reproduction of teleosts. The ovaries were processed by histological technique and stained with hematoxylin‐eosin (H‐E), Masson's trichrome, toluidine blue and periodic acid‐Schiff (PAS). The gonoduct is divided into three regions: cephalic, middle, and caudal. In the cephalic and middle regions there are mucosal folds that extend into the gonoductal lumen, forming structures similar to a cervix. The caudal region has two portions: the anterior contains a dorsal invagination and exocrine glands among columnar cells; the posterior has a ventral flexion and stratified epithelium with apical secretory cells. The morphology of this epithelium indicates two functions: (a) secretory by the apical columnar cells, and (b) protection through the stratification. Another peculiarity of the caudal region is that both ducts, reproductive and digestive, converge in a common cavity at their caudal ends, forming a cloacal region. The histology of the gonoduct indicates relevant functions including: (1) the control of the luminal diameter by the muscle and the presence of mucosal folds, like a cervix; (2) the relationship with the spermatozoa during insemination and storing them in mucosal folds; (3) the support of immunological processes; (4) secretory activities; (5) forming the duct during birth; and (6) possibly, acts as a barrier against parasite infestations.     

Genital system development of Williamia radiata (Gastropoda, Siphonariidae)

The adult anatomy and development of the genital system of the limpet Williamia radiata (Siphonariidae, Gastropoda) was investigated by means of 3D computer reconstruction and visualization of serial resin and paraplast light microscopical sections. As is characteristic for siphonariid species, the adult genital system consists of a single duct, the spermoviduct leading from the nidamental glandular system to the anteriorly located genital atrium with opening. The copulatory complex as well as the bursa copulatrix also open into the common genital atrium. The genital system develops from three separate anlagen. The posterior one appears first at a body length of 0.7 mm and gives rise to the ovotestis and part of the hermaphrodite duct. The nidamental glandular complex, the fertilization pouch-spermatheca complex, part of the hermaphrodite duct, the posterior part of the spermoviduct, and the bursa copulatrix develop later from the pallial anlage. Finally, the anterior anlage is formed on the right side of the head and gives rise to the genital atrium, the copulatory complex, and the anterior spermoviduct. This formation of the genital system from three, locally separated anlagen, differs strikingly from that of most other species of the Euthyneura. In both the Nudibranchia (Opisthobranchia) and the Stylommatophora (Pulmonata) development proceeds from a single site. We regarded this as a secondary condition as a result of derived features like heterochronies in development in these taxa. Comparison of development with that of other species of the Pulmonata allows conclusions on homology. The homology of the bursa copulatrix within the pulmonates is confirmed. The two separate chambers inside the spermoviduct of W. radiata correspond to oviduct and vas deferens of the freshwater Basommatophora.     

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.     

Hybridization between Helicoverpa armigera and Helicoverpa assulta (Lepidoptera: Noctuidae): development and morphological characterization of F1 hybrids

Reciprocal hybridizations between Helicoverpa armigera (Hübner) and Helicoverpa assulta (Guenée) were studied. The cross between females of H. armigera and males of H. assulta yielded only fertile males and sterile individuals lacking an aedeagus, valva or ostium bursae. A total of 492 larvae of the F1 generation were obtained and 374 of these completed larval development and pupated. Only 203 pupae were morphologically normal males, the remaining 171 pupae were malformed. Larvae and pupae that gave rise to morphologically abnormal adults exhibited longer development times. Sterility was not only associated with malformed external sex organs, but also a range of abnormalities of the internal reproductive system: (i) loss of internal reproductive organs, (ii) with one to three copies of an undeveloped bursa copulatrix; or (iii) with one or two undeveloped testes. Normal male hybrid adults showed higher flight activity in comparison with males of both species. In contrast, the cross between females of H. assulta and males of H. armigera yielded morphologically normal offspring (80 males and 83 females). The interaction of the Z-chromosome from H. assulta with autosomes from H. armigera might result in morphological abnormalities found in hybrids and backcrosses, and maternal-zygotic incompatibilities might contribute to sex bias attributed to hybrid inviability.     

长足大竹象生殖系统的形态解剖研究

解剖研究了长足大竹象雌雄虫牛殖系统的构造.该虫的雌性生殖系统包括一对卵巢、一对侧输卵管、中输卵管、交配囊、受精囊、生殖腔、产卵器;雄性生殖系统由一对睾九、一对输精管、一对附腺、射精管和交配器组成.     

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。     

Is Doris georgiensis García et al. 1993 a distinct species of dorid nudibranch?

The reproductive system and radular morphology of several specimens of Doris collected in the Antarctic Ocean were examined in an attempt to clarify the taxonomic status of Doris kerguelenensis and its possible synonym Doris georgiensis. A comparative analysis of the samples was conducted to find correlations between the anatomy of the bursa copulatrix and variations in the radular morphology and external features of these two species. Some consistent differences were found, suggesting the possibility of the existence of two distinct species, but further investigations are necessary to substantiate these conclusions.     

中国四川北部一沟颈螺新种(柄眼目,艾纳螺总科)描述(英文)

描述了栖息于甘肃南部的陆生贝类艾纳螺科1新种,南坪沟颈螺Holcauchen nanping sp. nov.。与各已知种相比,南坪沟颈螺的次体螺层最膨大;腭壁板齿及腔壁齿阙如;具1枚强壮轴唇齿;在生殖系统中,鞭状器小,乳突状;成荚器无盲囊;纳精囊管分支盲管缺乏。南坪沟颈螺,新种Holcauchen nanping sp. nov. (图1～4)鉴别特征次体螺层最膨大;无腭壁板齿与腔壁齿;轴唇齿1枚,强壮。鞭状器小乳突状;成荚器无盲囊;纳精囊管分支盲管缺乏。正模,HBUMM06584-specimen 1,具软体部的成体,四川省九寨沟县风成寺,2011-08-14,吴岷、徐沁、Prem B.Buhda采。模式标本保存于河北大学博物馆。词源:新种种名源自九寨沟县旧称"南坪"。     

Ultrastructure of the male reproductive system and of spermatogenesis in the viviparous brittle-star,Amphipholis squamata

The fine structure of the male reproductive system of the hermaphroditic brittle-star, Amphipholis squamata, has been studied in specimens from both the Pacific coast (Washington) and the Atlantic coast (New Hampshire). Each testis is a small (100-μm) sphere and is attached to the internal wall of the bursa by peritoneal suspensor cells. Occasional flagellated cells are found on the external surface of the testis. The testicular wall of A. squamata is a multilayered structure, similar to that of other ophiuroids, but the hemal sinus is PAS negative in this species. Germinal cells are surrounded throughout their development by the filopodia of interstitial cells. Adjacent interstitial cells are interconnected, and thus form a structural network within the testis. Positionally and functionally, the interstitial cells resemble Sertoli cells; however, their origin, behavior and ultrastructure are different in many ways. Spermatogenesis includes a series of cyclical changes (aspermatogenic phase, proliferative phase, differentiative phase, and evacuative phase). Within a single testis, the resulting production of sperm is in short pulses, but if all 10 testes are taken together, sperm are produced continuously throughout the year. The events of spermiogenesis closely follow those that have been described in other echinoderms. However, we have provided new information on the release of excess cell membranes and the fusion process of mitochondria. The mature spermatozoa of A. squamata are flagellated and motile, and have “primitive” structural features, in spite of the fact that they fertilize the eggs inside the genital bursae. The spermatozoa do not, as was previously thought, enter the bursa by rupture of the adjacent walls. Instead, they are ejaculated through a gonoduct into the rapid incurrent flow of water entering the bursa. The locomotion of the spermatozoa is in eccentric spirals, due to the unusually large angle at which the flagellum is inserted into the base of the sperm.     

Intraspecific transfer of cantharidin within selected members of the family Meloidae (Insecta: Coleoptera)

The use of deuterium-labelled cantharidin (CAN-D(2)) to study details of cantharidin transfer in blister beetles indicates that the dynamics of organ-selective cantharidin accumulation may differ over time. Although the accessory glands absorb a high amount of CAN-D(2) in the short term, they ultimately accumulate less than the testes. Confirming previous studies, the last steps in the pathway of biosynthesis of cantharidin occur in the male's body distantly from the reproductive system but the ultimate product, cantharidin, is transported into the male reproductive tract via the membrane of the accessory glands. From there it first transfers preferentially to the epididimis and the vas deferens, followed by final deposition in the testes. Most, if not all, of the cantharidin passes internally within the sexual organs; hemolymph transport is not involved. In female meloids, cantharidin enters the genitalia from the male as a nuptial gift. High amounts are first absorbed by the spermatophoral receptacle followed by spreading through the ovaries and an ultimate accumulation in the eggs. The amount taken up by the ovaries remains considerably lower than that of the receptacle. Over time these two organs stop accumulating cantharidin, while the bursa copulatrix starts to incorporate the gift actively. The accumulated amount taken up by bursa is mainly supplied by the receptacle and ovaries suggesting that an internal transfer of cantharidin is used in females as the main transport route.