Sperm transfer and reproductive biology in species of hermaphroditic bivalves (Galeommatoidea: Montacutidae)

The histology of the reproductive organs is studied in the protandric hermaphroditic Tellimya ferruginosa. In NW Europe the species reproduces from May through August. Sperm transfer takes place when mature testis follicles are transplanted to the gills or walls of the mantle cavity in recipient hermaphroditic or female bivalves. Transplantation is accompanied by histological changes and sperm cells are released when transplants perish with age. Details are given on the reproduction in Montacuta percompressa which takes place from March through October in North Carolina, USA. All shelled bivalves are females and it is postulated that spermatogenic bodies attached to gills or other surfaces in the female's mantle cavity and previously considered to be dwarf males arise from transplanted larval gonads. The ultrastructure of the euspermatozoa and/or the anucleate paraspermatozoa is described in T. ferruginosa, T. tenella, and M. percompressa. The sperm of the first two species share a number of significant apomorphies with those of another montacutid, Brachiomya stigmatica. In the simultaneous hermaphroditic M. substriata the nucleate paraspermatozoa associate with the euspermatozoa to form spherical spermatozeugmata that are stored in the testis.     

Morphology, structure of dimorphic sperm, and reproduction in the hermaphroditic commensal bivalve Pseudopythina tsurumaru (Galeommatoidea: Kellidae)

In Japan Pseudopythina tsurumaru is an up to 10.8 mm-long commensal of the burrowing sea cucumber Protankyra bidentata, whereas in Hong Kong the same species is smaller and associated with the crab Hexapus anfractus, itself a commensal of P. bidentata. Japanese P. tsurumaru is a hermaphrodite tending towards protogyny maturing to a female when > or = 7 mm, and entering the hermaphroditic condition when > or = 9 mm long. In addition to normal euspermatozoa, the species produces 30-32 microm long and 7 x 8 microm broad spindle-shaped paraspermatozoa provided with a conical acrosome, a nucleus, and a bundle of approximately 15-16 flagella issuing from the head region. Paired pouch-formed seminal receptacles normally occur in bivalves > or = 6 mm. Bulk sperm transfer presumably takes place by way of spermatozeugmata formed by the two types of sperm cells. Exogenous euspermatozoa attach to particular nonepithelial cells that occupy the interior of the receptacles. These cells, together with their associated sperm, are probably released as syncytial sperm-carrying bodies into the suprabranchial chamber, where the ova are fertilized.     

Sperm dimorphism and spermatozeugmata in the commensal bivalve Pseudopythina macrophthalmensis (Galeommatoidea, Kelliidae)

The bivalve Pseudopythina macrophthalmensis (Galeommatoidea) is a commensal with the crab Macrophthalmus convexus (Ocypodidae) in Okinawa Prefecture, Japan. It is a protandric hermaphrodite which incubates the 65-μm large ova in the suprabranchial cavity. The species produces two types of sperm, which were studied with the electron microscope. The euspermatozoon has an elongate 2.8-μm-long, pointed acrosome, a slender 12- to 13-μm-long nucleus and a middlepiece containing several closely packed mitochondria arranged as a 5.5- to 6.0-μm-long sheath around the basis of the flagellum. The paraspermato- zoon is vermiform, 220-μm-long and up to 5-μm-broad. Anteriorly there is a ca 7-μm-long bullet-shaped acrosome followed by a subcylindrical 3.0- to 4.7-μm-long nucleus. Adjacent to the nucleus occurs a bundle of 26–42 40-μm-long flagella. The cytoplasm is packed with spherical lipid droplets and ovoid granules of unknown composition. Sperm of both types aggregate to form spermatozeugmata, which were found in the posterior mantle cavity or in paired seminal receptacles. Within the receptacles the euspermatozoa dissociate themselves from the spermatozeugma and become attached to the epithelial lining of the receptacle whereas the paraspermatozoa presumably disintegrate. The possible significance of the two types of sperm is discussed in the light of their presumed functions in gastropods. Accepted: 9 November 2000     

Ultrastructure of the seminal receptacle and the dimorphic sperm in the commensal bivalve Mysella bidentata (Veneroida; Galeommatoidea; Montacutidae)

Jespersen, Å. and Lützen, J. 2001 . Ultrastructure of the seminal receptacle and the dimorphic sperm in the commensal bivalve Mysella bidentata (Veneroida: Galeommatoidea: Montacutidae). — Acta Zoologica (Stockholm) 82 : 107–115
The seminal receptacle and the euspermatozoa and paraspermatozoa of Mysella bidentata were examined at an ultrastructural level and the results were compared with earlier findings of the same and other species of the Montacutidae. The euspermatozoon has a slender 13 µm long nucleus and a 1.1 µm long bullet-shaped acrosome. The acrosome of the paraspermatozoon is almost identical in ultrastructure to that of the euspermatozoa but is longer (1.9 µm) and more slender and is bent at an angle to the diminutive nucleus (1.1 µm long). The unpaired seminal receptacle is lined by a heavily ciliated epithelium and a non-ciliated epithelium with short and broad microvilli. Euspermatozoa only are stored in the receptacle. They are densely packed and orientated with their heads towards the non-ciliated epithelium. In this position they develop numerous extremely fine microvilli from the acrosome which apparently serve to attach them to the epithelial microvillar surface. Stored sperm may presumably remain functional for at least six months. A possible function of paraspermatozoa could be to clump sperm into sperm bags to keep them in suspension.     

The reproductive system of Osedax (Annelida,Siboglinidae): ovary structure,sperm ultrastructure,and fertilization mode

Osedax is a genus of siboglinid annelids in which the females live on dead vertebrate bones on the seafloor. These females have a posterior end that lies within the bone and contains the ovarian tissue, as well as the “roots” involved with bone degradation and nutrition. The males are microscopic and live as “harems” in the lumen of the gelatinous tube that surrounds the female trunk, well away from the ovary. Females are known to spawn fertilized primary oocytes, suggesting internal fertilization. However, little is known about sperm transfer, sperm storage, or the location of fertilization, and the morphology of the female reproductive system has not been described and compared with the reproductive systems of other siboglinids. A 3D‐reconstruction of the ovisac of Osedax showed ovarian tissue with multiple lobes and mature oocytes stored in a “uterus” before being released through the single oviduct. The oviduct emerges as a gonopore on the trunk and travels along the trunk to finally open to the seawater as a thin cylindrical tube among the crown of palps. Light and transmission electron microscopy of mature Osedax sperm revealed elongate heads consisting of a nucleus with helical grooves occupied by mitochondria. In contrast to other Siboglinidae, Osedax sperm are not packaged into spermatophores or spermatozeugmata, and Osedax females lack a discrete region for sperm storage. Transmission electron microscopy and fluorescence microscopy allowed detection of sperm associated with ovarian tissue of the female ovisac of four different Osedax species. This provides the first evidence for the site of internal fertilization in Osedax. A heart body was found in the circulatory system, as seen in other siboglinids and some other annelids. The possible presence of nephridia in the anterior ovisac region was also documented. These morphological features provide new insights for comparing the regionalization of Osedax females in relation to other siboglinids.     

Reproductive morphology of Brittanichthys axelrodi (Teleostei: Characidae), a miniature inseminating fish from South America

Light and electron microscopy were used to investigate the morphology of reproductive characters in a characid fish, Brittanichthys axelrodi. Spermatozoa were found in ovaries of females, thereby confirming insemination in this species. Bony hooks can be found on the fourth unbranched ray and branched rays 1-4 of the anal fin and the unique sigmoidally-curved ray of the caudal fin in mature males. Testes have three distinct regions: an anterior spermatogenic region, an aspermatogenic middle region lined with a simple squamous epithelium and used for storage of mature spermatozoa, and a posterior region of coiled chambers lined with a high simple cuboidal epithelium. The most posterior region appears to be instrumental in the formation and storage of spermatozeugmata, unencapsulated sperm packets. Thus far, this tripartite testis morphology is unique among characids. The mature spermatozoon has an elongate nucleus ( approximately 5 microm in length). A striated rootlet originates at the anterior end of the distal centriole and continues to the anterior tip of the cell. The striated rootlet wraps around the entire ventral area of the anterior part of the nucleus and appears to continue around the anterior tip of the nucleus and down the dorsal side as electron-dense material. Several large, spherical mitochondria ( approximately 0.6 microm in diameter) with lamellar cristae overlap the posterior end of the nucleus and continue beyond together with the cytoplasmic collar that contains the flagellum which lacks axonemal fins. Each spermatozeugma is lanceolate in shape when sectioned mid-sagitally, with the core staining positively for mucopolysaccharides. In both sexes, the gonopore opens posterior to the anus, with the urinary pore having a separate opening posterior to the gonopore. Bands of skeletal muscle were found in the area of the male gonopore. These morphological features are likely linked to the reproductive mode of insemination, a trait that is so far as known, relatively rare among teleost fishes, but is proving increasingly frequent among certain groups of characid fishes.     

Ultrastructure of the annual cycle of female sperm storage in spermathecae of the torrent salamander, Rhyacotriton variegatus (Amphibia: Rhyacotritonidae)

This study is the first report on the ultrastructure of the sperm storage glands (spermathecae) in the salamander Rhyacotriton variegatus. The population studied is associated with cold-water, rocky streams of the redwood (Sequoia) zone in northern California. Males possess sperm in their vasa deferentia and undergo spermiation throughout the year, but mating is seasonal. Most females with large, vitellogenic follicles (2.0-3.9 mm mean dia.) collected from February-June contain sperm in their spermathecae, although some females with large follicles lack sperm. Other mature-size females collected during this period have small ovarian follicles (0.9-1.2 mm mean dia.) and lack stored sperm. All females collected from September-November have small follicles (0.6-1.6 mm mean dia.) and lack sperm, except in one instance in which a female collected in November had a small amount of degraded sperm, apparently retained from the previous breeding season. The spermathecae consist of simple tubulo-alveolar glands in which the neck tubules produce a mucoid secretory product, and the distal bulbs, where sperm are stored, contain secretory vacuoles of uniform density that stain positively for glycosaminoglycans. In specimens containing sperm, some bulbs have abundant sperm and others lack sperm, but the ultrastructure is similar in both conditions. The acini contain columnar epithelial cells with wide intercellular canaliculi, and a merocrine process releases the secretion. Spermiophagy occurs. In specimens from spring and summer with small ovarian follicles, the neck tubules are similar to those of breeding females, but the distal bulbs are reduced to cords of cells lacking a discernible lumen. Secretory activity in the distal bulbs is initiated in the fall. Spermathecae of R. variegatus are most similar to those of a stream-dwelling plethodontid, Eurycea cirrigera.     

Ultrastructure of spermiogenesis, sperm, and the spermatheca in Terebrasabella heterouncinata (Polychaeta: Sabellidae: Sabellinae)

Abstract. The sabellid polychaete, Terebrasabella heterouncinata, forms burrows in gastropod shells. It is a small, intratubular brooder that breeds semi‐continuously. It has been shown to self‐fertilize, but its reproductive biology suggests that some form of sperm transfer must occur between individuals. To gain an understanding of its fertilization biology, the ultrastructure of spermiogenesis and the sperm in T. heterouncinata was described, and the animal examined for sperm storage structures. Spermiogenesis occurs in clusters of eight spermatids. The mature sperm has an elongate nucleus and a bilaterally symmetrical acrosome with twisted subacrosomal spaces. The midpiece is short, with three crescent‐shaped mitochondria, and forms a tight sheath around the axoneme. A single spermatheca, which opens on the inner ventral part of the crown near the buccal region, is present. It is a simple blind‐ending duct that runs below the ventral nerve cord and is longer than 100 μm. This is the first record of a single spermatheca in Sabellidae. The shape of the sperm and the presence of a spermatheca confirm that individuals of T. heterouncinata produce ent‐aquasperm and would normally cross‐fertilize.     

Mechanisms of sperm removal and sperm transfer in Orthetrum coerulescens (Fabricius) (Odonata: Libellulidae)

ABSTRACT An account of the structure of the secondary genitalia of male Orthetrum coerulescens (Fabricius) is given together with an outline of the internal genitalia of the female. Observations on the experimental inflation of the penis are described. No sperm was released during fast inflations and deflations of the penis, but sustained inflation of the penis led to the release of a clear fluid followed by the slow discharge of sperm. Examination of natural copulations in the field has shown that rapid rhythmic movements at up to 5 Hz take place in the male's third abdominal segment throughout most of copulation, but towards the end they cease though inflation is maintained. It is suggested that the rapid movements coincide with the removal of rival sperm from the female and that sperm is transferred to the female only during the maintained inflation. Possible mechanisms of sperm translocation, sperm removal and sperm transfer are discussed with reference to male and female genital structures and the action of relevant muscles.     

Reproductive apparatus and male accessory structures in two batrachoid species (Teleostei, Batrachoididae)

The morphology of the genital apparatus of two batrachoid species, Opsanus tau and Porichthys notatus , was studied. The anatomical organization of the female reproductive apparatus is similar in both species but differences are observed in the rhythm of gametogenesis with individual oocyte production asynchronous in O. tau and group synchronous in P. notatus. The male reproductive apparatus is similarly organized in the two species, with both showing enlongated testes with an efferent duct system, two main testicular ducts, a common sperm duct, and a pair of multi-channel accessory structures. The sperm transport system consisting of the efferent duct system, main testicular ducts, and sperm duct is more developed in P. notatus and only in this species does it secrete sialoglycoproteins. Male accessory organs also secrete sialoglycoprotein in both species, but they appear more developed in O. tau. Intraspecific variability in development of accessory structures and mucin secretion was also observed between the two male morphs of P. notatus. Type I males, which build nests and perform parental care to eggs and fry, have larger accessory organs and more abundant secretion than type II males, which adopt opportunistic spawning tactics. The possible role played by mucins as components of the seminal fluid, in both species, and their inter-and intraspecific variability are discussed in the light of the reproductive biology and the presence of alternative male mating tactics.     

Reproduction and sperm structure in Galeommatidae (Bivalvia, Galeommatoidea)

Reproduction and/or sperm structure was studied in 18 species belonging to five genera of Galeommatidae from Phuket Island, Andaman Sea, Thailand, and Hong Kong. Incubation of the ova occurs in the inner and outer demibranchs of both gills, and sperm is most probably transferred to the ctenidial brood chamber as masses of agglutinated and non-encapsulated spermatozoa. The smallest specimens are males, but change into the female sex and there is a strong indication that some of the species are alternate hermaphrodites. Dwarf males occur in one species (Galeomma layardi). The sperm of all 18 species studied are of the ent-aquasperm type with a more or less hemispherical acrosome that is tilted with respect to the long axis of the moderately elongated nucleus. It is suggested that this type of sperm affords the best autapomorphy for the family Galeommatidae.     

The sperm structure of Cryptocercus punctulatus Scudder (Blattodea) and sperm evolution in Dictyoptera

Sperm of the dictyopteran key taxon Cryptocercus punctulatus was examined. It has largely maintained a blattodean groundplan condition, with a three‐layered acrosome, an elongate nucleus, a single centriole, a conspicuous centriole adjunct material, two connecting bands (=accessory bodies), and a long functional flagellum with a 9+9+2 axoneme provided with accessory tubules with 16 protofilaments and intertubular material. These sperm characters are shared with several other polyneopterans. The sperm of C. punctulatus is very similar to what is found in Periplaneta americana and species of other groups of roaches, including the sperm of Loboptera decipiens described here for the first time. The general sperm organization here described can be assumed for the groundplan of Insecta and Pterygota. The following evolutionary path can be suggested: after the split between Cryptocercidae and the common ancestor of Isoptera, the typical pattern of sperm formation was altered very distinctly, resulting in a duplication or multiplication (Mastotermitidae) of the centrioles. Mastotermes has maintained a certain sperm motility, but with a very unusual apparatus of multiple flagella with a 9+0 axoneme pattern. After the split into Mastotermitidae and the remaining Isoptera, sperm motility was completely abandoned, and different modifications of sperm components occurred, and even the loss of the sperm flagellum. J. Morphol. 276:361–369, 2015. © 2014 Wiley Periodicals, Inc.     

Remating, sperm transfer, and sperm displacement in the cactophilic species Drosophila buzzatii Patterson & Wheeler (Diptera: Drosophilidae)

The mating system of Drosophila buzzatii is characterized by short copulation duration, frequent remating in both males and females, and male ejaculate partitioning. Additional features of the system are strong sperm displacement and a high frequency of sterile matings. Remating frequencies and the effects of remating on various mating parameters were studied. In order to characterize variation, five isofemale lines from geographically distant localities in Australia (three localities), Brazil and the Canary Islands were used. Mating parameters studied were: premating time, copulation duration, interval between successive matings, and progeny number as a measure of sperm transfer. Variation for sperm displacement was studied in crosses between laboratory stocks and a number of isofemale lines from Australia. There were significant between‐line differences in female remating frequencies, premating time, copulation duration, interval between successive matings, and progeny numbers, indicating genetic variation for these traits. Females from the five lines mated on average 1.6 to 3.1 times in 4 h, with a maximum of eight matings for one female. The males were given a maximum of ten virgin females in sequence and more than one‐third of the males mated all ten females in the 2 h observation period. Copulation duration decreased and interval between matings increased with copulation number in multiply mated males. Mean copulation duration was c. 2 min. Sperm transfer, measured as the average number of progeny from a single mating, was low (c. 25) and multiply mated females gave more progeny than single mated females, although with much lower progeny numbers than observed in wild‐caught non‐virgin females. A surprisingly high proportion of observed matings gave no progeny, i.e. they were sterile matings. Sperm displacement was strong in most crosses and remained strong in multiply mated females. The results are discussed in relation to the evolution of mating patterns in Drosophila.     

Sperm ultrastructure in Kellia suborbicularis (Bivalvia: Galeommatoidea: Kellidae)

The sperm cells of Kellia suborbicularis are narrow with a short bullet‐shaped acrosome, a 5.0–5.5 µm long and 0.4–0.6 µm broad nucleus, and a short midpiece with a ring of five mitochondria. The disposition of the subacrosomal substance into a coronet‐like formation is unique, and the sperm structure offers no clue to the relationship between Kellia and other galeommatoidean genera. The possible significance of narrow elongate sperm for their entry into the brood pouch is discussed.     

Sperm and spermatozeugma ultrastructure in the inseminating species Tyttocharax cochui, T. tambopatensis, and Scopaeocharax rhinodus (Pisces: Teleostei: Characidae: Glandulocaudinae: Xenurobryconini)

This article presents the scanning and transmission electron microscopy of the spermatozoa and sperm packets of three inseminating species of the glandulocaudine tribe Xenurobryconini. All three species, Scopaeocharax rhinodus, Tyttocharax cochui, and T. tambopatensis produce unencapsulated sperm packets (= spermatozeugmata) of similar morphology. The external anterior surface of each spermatozeugma is comprised of elongate sperm heads arranged in parallel, and the posterior part is made up of tightly packed flagella. The interior of the anterior portion consists of alternating layers of sperm heads and flagella. The remarkable integrity of each packet appears to be maintained through an electron-dense secretion seen among all parts of the cells. Spermatozeugma formation takes place within the spermatocysts at the end of spermiogenesis and at spermiation fully formed packets are released. Morphology of the mature spermatozoa was similar in all three species. Each nucleus is elongate, flattened along most of its length, and tapers at either end. The two centrioles are nearly parallel to one another and are located just anterior to the nucleus. Elongate mitochondria are located along the nucleus. The single flagellum, which lacks axonemal fins, is initially contained within a short cytoplasmic collar. Accessory microtubules run parallel to the long axis of the nucleus just beneath the plasma membrane. During spermiogenesis, no nuclear rotation occurs and the cytoplasmic canal containing the flagellum elongates along with the nucleus. However, prior to spermiation all but the anterior portion of the collar degenerates. The sperm modifications observed in these species are discussed as adaptations to the unique reproductive habit of insemination.     

Numbers and size of sperm storage tubules and the duration of sperm storage in birds: a comparative study

Estimates of the numbers of sperm storage tubules (SSTs) in the utero-vaginal junction of 11 bird species are presented. Numbers of SSTs varied by a factor of 40 between species, and ranged from 500 to 20000. Body mass accounted for over 50% of the variation in SST mumbers. SST length was positively correlated with the length of spermatozoa across species. The duration of sperm storage was not correlated with the number of SSTs or the volume of sperm storage tissue. However, the number of 'active' SSTs appears to vary between species and it was not possible to make allowance for this. Sperm storage duration was weakly, positively correlated wth clutch size, but showed a significant positive relationship with the number of days over which laying occurred. The number of SSTs was also positively correlated with the number of sperm per ejaculate. The best predictor of sperm storage duration was a multiple regression equation using the spread of laying and the length of sperm storage tubules. The duration of sperm storage in birds which remain together during the pre-laying period is such that a single insemination immediately before the start of laying could fertilize the entire clutch.     

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.