Ultrastructure of spermiogenesis and synthesis of enigmatic cytoplasmic inclusions in the spirally shaped spermatozoon of the polychaete Spio setosa (Annelida: Spionidae)

The sperm of Spio setosa (Polychaeta, Spionidae) are known to be very unusual in form; here, spermiogenesis and the structure of the spermatozoon in this species are described by transmission electron microscopy. While spermiogenesis is similar to that described for many other polychaetes, two notable exceptions to this process include the synthesis of abundant ring‐shaped and tubular, membrane‐bounded cytoplasmic inclusions in the midpiece, and the differentiation of a spirally shaped sperm head. Spermatids develop as free‐floating tetrads in the male's coelom. A microtubular manchette does not develop during chromatin condensation and nuclear elongation, and the spiral acrosome forms as a single Golgi‐derived vesicle that migrates anteriorly to become housed in a deep anterior nuclear fossa. Early in spermiogenesis, numerous Golgi‐derived, membrane‐bounded cytoplasmic inclusions appear in the cytoplasm; these ultimately occupy the sperm midpiece only. The mature spermatozoon in the male has a 15‐μm‐long head consisting of a nucleus coiled like a spring and a spiral acrosome with differentiated substructure, the posterior two thirds of which sits in an anterior nuclear fossa. The midpiece is wider than the rest of the spermatozoon and contains 9–10 spherical mitochondria surrounding the two centrioles, as well as numerous membrane‐bounded conoid and tubular cytoplasmic inclusions. The axoneme has a 9 + 2 arrangement of microtubules. By contrast, stored sperm in the female's seminal receptacles have lost the midpiece inclusions but contain an abundance of glycogen. The function of the midpiece inclusions remains unresolved, and the significance of their absence in stored sperm within the seminal receptacle and the appearance of midpiece glycogen stores remains unclear and requires additional investigation.     

Ultrastructural study of spermiogenesis and the testicular and spermathecal spermatozoon of the Gonochoristic Tardigrade Xerobiotus pseudohufelandi (Eutardigrada,Macrobiotidae)

This paper investigates by scanning and transmission electron microscopy spermiogenesis and spermatozoon morphology of the gonochoristic eutardigrade Xerobiotus pseudohufelandi (Macrobiotidae). During spermiogenesis clusters of spermatids are connected by cytoplasmic bridges that persist up to an advanced stage of maturation. Spermiogenesis is characterized by distinctive modifications of the nucleus and by the differentiation of an acrosome, tail and substantial midpiece. Testicular spermatozoa are folded with the hinge located between the head and midpiece, thus resembling a nut-cracker. The head includes a rod-shaped, bilayered acrosome and an elongated, helicoidal nucleus with condensed chromatin. The large kidney-shaped midpiece has hemispherical swellings/ovoid elements surrounding the centriole and an incomplete mitochondrial sleeve. The flagellum contains a ‘9+2’ axoneme and terminates in a tuft of microtubules. Spermathecal spermatozoa always have linear profiles. The acrosome and nucleus have the same morphological pattern as in testicular spermatozoa, whereas the midpiece is thin and lacks the hemispherical swellings, and the tail is reduced to a short stub. Functional considerations are presented, based upon this different morphology. Moreover, phyletic comparisons are made on the basis of sperm morphology, both for the family Macrobiotidae and the class Eutardigrada. J. Morphol. 234:11–24, 1997. © 1997 Wiley-Liss, Inc.     

The ultrastructure of amberjack (Seriola dumerilii) sperm

Scanning and transmission electron microscopy were used to investigate the fine structure of sperm of the Mediterranean amberjack Seriola dumerilii. Each spermatozoon has an ovoid head which lacks an acrosome, a short, irregularly-shaped midpiece and a long flagellar tail. The midpiece houses eight spherical mitochondria, which are separated from the axoneme by the cytoplasmic canal. The centrioles are arranged approximately at right angles to each other. The proximal centriole lies inside, and the distal centriole outside, the nuclear fossa. The flagellum is inserted eccentrically into the head and is tangential to the nucleus, so that the spermatozoon is asymmetrical. It contains the conventional 9 + 2 axoneme, shows intratubular differentiations in the A microtubules of doublets 1, 2, 5 and 6, and possesses one pair of lateral fins. On the basis of its ultrastructural organization, the amberjack sperm resembles type II sperm as defined previously, except for the presence of the proximal centriole inside the nuclear fossa. This could result from a partial rotation of the nucleus during spermiogenesis.     

Ultrastructure of spermiogenesis in a freshwater stingray, Himantura signifer

Ultrastructural changes of spermatids during spermiogenesis in a freshwater stingray, Himantura signifer, are described. Differentiation of spermatids begins with modification of the nuclear envelope adjacent to the Golgi apparatus, before the attachment of the acrosomal vesicle. A fibrous nuclear sheath extends over the nuclear surface from the site of acrosomal adherence. The conical apical acrosome is formed during nuclear elongation. At the same time, chromatin fibers shift from an initially random arrangement, assume a longitudinal orientation, and become helical before final nuclear condensation. An axial midpiece rod is formed at the posterior end of nucleus and connects to the base of the sperm tail. Numerous spherical mitochondria surround the midpiece axis. The tail originating from the posterior end of the midpiece is composed of the usual 9 + 2 axoneme accompanied by two longitudinal columns, which are equal in size and round in cross section. The two longitudinal columns are absent at the end piece. A distinctive feature of freshwater stingray sperm is its spiral configuration.     

Fine structure of the spermatozoa of Crassostrea gigas (Mollusca,Bivalvia)

We describe sperm ultrastructure and acrosome differentiation during spermiogenesis in Crassostrea gigas (Mollusca Bivalvia). The sperm cell is a uniflagellated cell of the primitive type. The head region contains a rounded or conical nucleus surmounted by small acrosome. This organelle consists of a membrane-bound acrosomal granule, the contents of which have a homogeneous density, except in the anterior region, which is positive for PTA. The acrosome also surrounds the perforatorium, which includes oriented fibrillar elements: this is the axial body. The middle piece contains four mitochondria encircling two perpendicular centrioles. The distal centriole is provided with a system of mechanical fixation to the plasma membrane, consisting of nine fibers in radial arrangement. The tail flagellum, about 50 m?m long, contains the usual microtubular axoneme. © 1993 Wiley-Liss, Inc.     

Four lines of spermatid development and dimorphic spermatozoa in the sea urchin Anthocidaris crassispina (Echinodermata, Echinoida)

 The process of sperm development in the sea urchin Anthocidaris crassispina was studied by light and electron microscopy. Similar to other echinoids studied, a single flagellum, striated rootlet and nuage-like materials were present in spermatogonia of A. crassispina. Spermatocytes near the diplotene stage showed intracellular localization of the axoneme which appeared to be a retracted flagellum prior to cell division. Fibrous filaments were associated with a proximal centriole in spermatocytes and spermatids and might be involved in movement of the proximal centriole. An acrosomal vesicle was developed and a residual body was formed in spermatids. The special development patterns in A. crassispina attributed to the presence of two patterns of tail development and two patterns of mitochondrial development during spermiogenesis. These four lines of spermiogenesis resulted in the formation of four morphological types of sperm cell, i.e. sperms with: (1) a symmetrical midpiece and posterior tail, (2) an asymmetrical midpiece and posterior tail, (3) a symmetrical midpiece and bent tail and (4) an asymmetrical midpiece and bent tail. Sperm cells with bent tails (type 3+4) were probably still at the late spermatid stage because results of scanning electron microscopy demonstrated gradual detachment and eventual straightening of the bent tail, and their percentage occurrence in the sperm population decreased significantly (P<0.05) towards the spawning season of A. crassispina. Spermatozoa with a symmetrical midpiece were dominant (averaging 70% occurrence in the sperm population) over those with an asymmetrical midpiece. The dimorphic spermatozoa in A. crassispina (types 1, 2) are both considered to be euspermatozoa as their morphology is typical for Echinoida. Accepted: 4 May 1998     

The ultrastructure of spermatozoa and spermiogenesis in pyramidellid gastropods,and its systematic importance

Ultrastructural observations on spermiogenesis and spermatozoa of selected pyramidellid gastropods (species ofTurbonilla, Pyrgulina, Cingulina andHinemoa) are presented. During spermatid developement, the condensing nucleus becomes initially anterio-posteriorly compressed or sometimes cup-shaped. Concurrently, the acrosomal complex attaches to an electrondense layer at the presumptive anterior pole of the nucleus, while at the opposite (posterior) pole of the nucleus a shallow invagination is formed to accommodate the centriolar derivative. Midpiece formation begins soon after these events have taken place, and involves the following processes: (1) the wrapping of individual mitochondria around the axoneme/coarse fibre complex; (2) later internal metamorphosis resulting in replacement of cristae by paracrystalline layers which envelope the matrix material; and (3) formation of a glycogen-filled helix within the mitochondrial derivative (via a secondary wrapping of mitochondria). Advanced stages of nuclear condensation (elongation, transformation of fibres into lamellae, subsequent compaction) and midpiece formation proceed within a microtubular sheath (‘manchette’). Pyramidellid spermatozoa consist of an acrosomal complex (round to ovoid apical vesicle; column-shaped acrosomal pedestal), helically-keeled nucleus (short, 7–10 μm long, shallow basal invagination for axoneme/coarse fibre attachment), elongate helical midpiece (composed of axoneme, coarse fibres, paracrystalline and matrix materials, glycogen-filled helix), glycogen piece (length variable, preceeded by a dense ring structure at junction with midpiece). The features of developing and mature spermatozoa observed in the Pyramidellidae are as observed in opisthobranch and pulmonate gastropods indicating that the Pyramidelloidea should be placed within the Euthyneura/Heterobranchia, most appropriately as a member group of the Opisthobranchia.     

Fine structure of the spermatozoon of Marthasterias glacialis (Linnaeus) (Echinodermata; Asteroidea), with special reference to acrosomal morphology

The sperm of Marthasterias glacialis (Linnaeus) was studied by light and electron microscopy. It is a long uniflagellated cell of the “primitive” type. The head has a spherical shape and contains a nucleus with a spheroid acrosome lying in a cup-shaped anterior fossa. The acrosome is formed by an acrosomal vesicle surrounded by the periacrosomal material. The basal specializations of the acrosomal vesicle show a clear differentiation of its constituents resembling the structure of membrane. The midpiece contains a very large annular mitochondrion which encircles two perpendicular centrioles. The distal centriole is in close association with a pericentriolar radial complex. The tail, containing a common microtubular axoneme, is projected to a variable position.     

Spermatozoa and sperm packages of the European troglophylous scorpion Belisarius xambeui Simon, 1879 (Troglotayosicidae, Scorpiones)

Studies on the sperm morphology in scorpions are rare, but the existing investigations already revealed a remarkable interfamiliar diversity. The present study reports for the first time on the spermatozoa and sperm packages of a representative of the family Troglotayosicidae, the troglophylous species Belisarius xambeui. The spermatozoa are characterized by (1) a thread-like nucleus, which is slightly bent anteriorly; (2) an asymmetrical cap-like acrosomal vacuole, which encloses the anterior tip of the nucleus; an acrosomal filament is absent; (3) an axoneme with a 9 + 0 microtubular pattern; (4) a midpiece consisting of elongated mitochondria coiling around the axoneme; the number can vary between 3 and 6 (mostly 4). At the end of spermiogenesis, the spermatozoa aggregate in order to form oval-shaped sperm packages in which all sperm cells show the same orientation. A single package consists of approximately 150 sperms. A secretion sheath is always absent. The present results might provide new characters for further systematic studies and their phylogenetic implications are briefly discussed.     

The spermatozoon of Oikopleura dioica Fol (Larvacea,Tunicata)

Summary The spermatozoon of Oikopleura dioica is about 30 m long, with a spherical head, about 1 m wide, a 3 m long and 1 m wide midpiece, and a 25 m long tail with a tapered end piece. The head contains a nucleus with the chromatin volume limited to about 0.1 m³. A small acrosome is found in an anterior inpocketing, and a flagellar basal body in a posterior inpocketing of the nucleus. The midpiece contains a single mitochondrion with the flagellar axoneme embedded in a groove along its medial surface. The flagellar axoneme has the typical 9 + 2 substructure, and the basal body the typical 9+0 substructure. A second centriole and special anchoring fibres are absent.     

Fine structure of acrosome biogenesis and of mature sperm in the bivalve molluscs<Emphasis Type="Italic"> Glycymeris</Emphasis> sp. (Pteriomorphia) and<Emphasis Type="Italic"> Eurhomalea rufa</Emphasis> (Heterodonta)

Proacrosomal vesicles form during the pachytene stage, being synthetized by the Golgi complex in Glycymeris sp., and by both the Golgi and the rough endoplasmic reticulum in Eurhomalea rufa. During early spermiogenesis, a single acrosomal vesicle forms and its apex becomes linked to the plasma membrane while it migrates. In Glycymeris sp., the acrosomal vesicle then turns cap-shaped (1.8 μm) and acquires a complex substructure. In E. rufa, proacrosomal vesicles differentiate their contents while still at the premeiotic stage; as the acrosomal vesicle matures and its contents further differentiate, it elongates and becomes longer than the nucleus (3.2 μm), while the subacrosomal space develops a perforatorium. Before condensation, chromatin turns fibrillar in Glycymeris sp., whereas it acquires a cordonal pattern in E. rufa. Accordingly, the sperm nucleus of Glycymeris sp. is conical and elongated (8.3 μm), and that of E. rufa is short and ovoid (1.1 μm). In the midpiece (Glycymeris sp.: 1.1 μm; E. rufa: 0.8 μm), both species have four mitochondria encircling two linked orthogonal (Glycymeris sp.) or orthogonal and tilted (30–40°; E. rufa) centrioles. In comparison with other Arcoida species, sperm of Glycymeris sp. appear distinct due to the presence of an elongated nucleus, a highly differentiated acrosome, and four instead of five mitochondria. The same occurs with E. rufa regarding other Veneracea species, with the acrosome of the mature sperm strongly resembling that of the recent Mytilinae. Electronic Publication     

Fine structure of spermatozoa in the blackspot sea bream Pagellus bogaraveo (Brünnich, 1768) with some considerations about the centriolar complex

Scanning and transmission electron microscopy were used to investigate the fine structure of the sperm of the sparid fish Pagellus bogaraveo.The spermatozoon of P. bogaraveo belongs, like that of the other sparid fish, to the teleostean “type I” spermatozoon with the flagellar axis insert perpendicular to the nuclear fossa. It has an ovoidal head, a short, cylindrically shaped midpiece and a long tail region. The nucleus reveals a deep invagination (nuclear fossa), in which the centriolar complex is located, and a satellite nuclear notch shaped like a golf club. The two centrioles are perpendicular to each other and show a conventional “9+0” pattern. The distal centriole is attached to the nuclear envelope by means of basal feet and radial fibers made of electron-dense material. Below the basal plate, plasma membrane pinches in, and the necklace, a specialized connection joining axonemal doublets to the plasma membrane, is visible. The short midpiece houses one mitochondrion. The flagellum is perpendicularly and eccentrically with respect to the nucleus and contains the conventional “9+2” axoneme.     

Ultrastructure of Thunnus thynnus and Euthynnus allettevatus spermatozoa

The spermatozoa of Thunnus thynnus and Euthynnus alletteratus consist of an acrosome-less head (comprising the ovoid nucleus and the short midpiece) and a long flagellar tail that contains the conventional 9 + 2 axoneme and lacks lateral fins. The centrioles are arranged at approximately right angles and lie outside of a shallow nuclear groove. The flagellum inserts laterally on the nucleus, therefore the spermatozoon is asymmetrical. The midpiece contains a few mitochondria which are separated from the axoneme by the cytoplasmic canal; they are spherical in T. thynnus and elongate, somewhat irregular in E. alletteratus . Although the main ultrastructural characteristics of the spermatozoa appear to indicate a great homogeneity in the sperm morphology within the family Scombridae, small species-specific divergences may be of use in systematics.     

Sperm ultrastructure and a new type of spermiogenesis in two species of Pimelodidae, with a comparative review of sperm ultrastructure in Siluriformes (Teleostei: Ostariophysi)

During spermiogenesis, the spermatids of the pimelodid species Pimelodus maculatus and Pseudoplatystoma fasciatum show a central flagellum development, no rotation of the nucleus, and no nuclear fossa formation, in contrast to all previously described spermatids of Teleostei. These characteristics are interpreted as belonging to a new type of spermiogenesis, named here type III, which is peculiar to the family Pimelodidae. In P. maculatus and P. fasciatum, spermatozoa possess a spherical head and no acrosome; their nucleus contains highly condensed, homogeneous chromatin with small electron-lucent areas; and a nuclear fossa is not present. The centriolar complex lies close to the nucleus. The midpiece is small, has no true cytoplasmic channel, and contains many elongate and interconnected vesicles. Several spherical to oblong mitochondria are located around the centriolar complex. The flagellum displays the classical axoneme (9+2) and no lateral fins. Only minor differences were observed among the pimelodid species and genera. Otherwise, spermiogenesis and spermatozoa in the two species of Pimelodidae studied exhibit many characteristics that are not found in other siluriform families, mainly the type III spermiogenesis.     

Ultrastructure of euspermatozoa and paraspermatozoa in the volutid snail <Emphasis Type="Italic">Adelomelon ancilla</Emphasis> (Mollusca: Caenogastropoda)

The ultrastructure of the euspermatozoa and the paraspermatozoa is investigated in Adelomelon ancilla, through histological section observed by transmission electron microscopy. Euspermatozoa of A. ancilla consists of: (1) a conical acrosomal vesicle (with a short basal invagination, constricted anteriorly) which is flattened at the apex and associated with an axial rod, a centrally perforated basal plate and a short accessory membrane, (2) a rod-shaped, solid and highly electron-dense nucleus (with a short basal fossa containing a centriolar complex and a initial portion of a 9 + 2 axoneme), (3) an elongate midpiece consisting of the axoneme sheathed by 5–6 helical mitochondrial elements each exhibiting a dense U-shaped outer layer, (4) an elongate glycogen piece (where the axoneme is sheathed by nine tracts of glycogen granules), (5) a dense annulus at the junction of the midpiece and glycogen piece, and (6) a short free tail region (where the axoneme is surrounded only by plasma membrane). We observed a parasperm in A. ancilla. This is vermiform in shape and is composed of multiple axonemes and extensive cytoplasm with numerous vesicles, and mitochondria are scattered inside the axonemes. Sperm of A. ancilla is characterized by the euspermatozoa type 2 and the paraspermatozoa morphology belongs to type 5. The U shaped electrodense mitochondrial element in the midpiece of the eusperm and the constriction in the acrosomal vesicle present in A. ancilla are exclusive. We suggest that these characteristics could have taxonomic importance, because these was observed in other volutids and have not been observed in the rest of caenogastropods studies. We consider that the morphology of paraspermatozoa in A. ancilla corresponds to the “lancet” type.     

Ultrastructure and potential taxonomic importance of euspermatozoa and paraspermatozoa in the volutid gastropods Zidona dufresnei and Provocator mirabilis (Caenogastropoda, Mollusca)

The ultrastructure of mature spermatozoa is investigated for the first time in the Volutidae, based on the commercially significant South American species Zidona dufresnei (Donovan, 1823) (fresh material) and supplemented with observations on testicular (museum) material of the deep sea New Zealand species Provocator mirabilis (Finlay, 1926). Euspermatozoa of Z. dufresnei (ex sperm duct) consist of: (1) a tall-conical acrosomal vesicle (with short basal invagination, constricted anteriorly) which is flattened anteriorly and associated with an axial rod, centrally perforate basal plate and short accessory membrane; (2) a rod-shaped, solid and highly electron-dense nucleus (with short basal fossa containing centriolar complex and initial portion of a 9 + 2 axoneme); (3) an elongate midpiece consisting of the axoneme sheathed by 5–6 helical mitochondrial elements, each exhibiting a dense U-shaped outer layer; (4) an elongate glycogen piece (axoneme sheathed by nine tracts of putative glycogen granules); (5) a dense annulus at the junction of the midpiece and glycogen piece and (6) a short free tail region (axoneme surrounded only by plasma membrane). Paraspermatozoa of Z. dufresnei are vermiform and dimorphic: the first type contains approximately 14–20 axonemes (arranged peripherally and interspersed with microtubules) and numerous oblong dense vesicles, numerous less dense (round) vesicles, occasional, large lipid-like vesicles, and scattered mitochondria; the second type contains 25–31 axonemes (peripherally arranged, interspersed with microtubules), occasional mitochondria and extensive cytoplasm. Results obtained for P. mirabilis from testis material are essentially as observed in Z. dufresnei, although the euspermatozoan acrosome still has to achieve its compressed transverse profile. Observations on paraspermatozoa were limited by fixation quality of available (testis) tissues, but these cells are similar to the first type of Zidona paraspermatozoa. Although most of the euspermatozoal features are also observed in many neotaenioglossans and neogastropods, the U-shaped outer layer of each mitochondrial element has not previously been reported and may prove a diagnostic feature of the Volutidae, the subfamily Zidoniinae or possibly only the Zidonini (in which Z. dufresnei and P. mirabilis are currently placed).     

Ultrastructural aspects of spermatogenesis in Calyptogena pacifica Dall 1891 (Vesicomyidae; Bivalvia)

The process of spermatogenesis and spermatozoon morphology was characterized from a deep‐sea bivalve, Calyptogena pacifica (Vesicomyidae, Pliocardiinae), a member of the superfamily Glossoidea, using light and electron microscopy. Spermatogenesis in C. pacifica is generally similar to that in shallow‐water bivalves but, the development of spermatogenic cells in this species has also some distinguishing features. First proacrosomal vesicles are observed in early spermatocytes I. Although, early appearance of proacrosomal vesicles is well known for bivalves, in C. pacifica, these vesicles are associated with electron‐dense material, which is located outside the limiting membrane of the proacrosomal vesicles and disappears in late spermatids. Another feature of spermatogenesis in C. pacifica is the localization of the axoneme and flagellum development. Early spermatogenic cells lack typical flagellum, while in spermatogonia, spermatocytes, and early spermatids, the axoneme is observed in the cytoplasm. In late spermatids, the axoneme is located along the nucleus, and the flagellum is oriented anteriorly. During sperm maturation, the bent flagellum is transformed into the typical posteriorly oriented tail. Spermatozoa of C. pacifica are of ect‐aqua sperm type with a bullet‐like head of about 5.8 μm in length and 1.8 μm in width, consisting of a well‐developed dome‐shaped acrosomal complex, an elongated barrel‐shaped nucleus filled with granular chromatin, and a midpiece with mainly four rounded mitochondria. A comparative analysis has shown a number of common traits in C. pacifica and Neotrapezium sublaevigatum.     

Ultrastructure of the sperm and spermatogenesis and spermiogenesis of Dina lineata (hirudinea,erpobdellidae)

The mature sperm of Dina lineata is of the modified type. The sperm are 48 μm long and 0.3 μm wide. The sperm are filiform and helicoidal cells with a distinct head, a midpiece, and a tail. There are two distinct regions in the head: the acrosome and the posterior acrosome, each with its own characteristic morphology. The midpiece is the mitochondrial region and has a single mitochondrion. Two distinct portions can be observed in the tail: the axonematic region and the terminal piece. In the process of spermatogenesis the early spermatogonia divide to form a poliplast of 512 spermatic cells. In the spermiogenesis the following sequential stages can be distinguished: elongation of the flagellum; reciprocal migration of mitochondria and Golgi complex; condensation of chromatin and formation of the posterior acrosome; spiralization of nuclear and mitochondrial regions; and, finally, formation of the anterior acrosome. The extreme morphological complexity of the Dina spermatozoon is related to the peculiar hypodermal fertilization which characterizes the erpobdellid family. Correlation between sperm morphology and fertilization biology in the Annelida is revised.     

Ultrastructure of sperm and male reproductive system in Lineus viridis (Heteronemertea, Nemertea)

Nemerteans possess serially arranged gonads that lie between the midgut pouches. In both sexes the gonads are lined with an epithelium. During maturity, they gain contact to the exterior by a ciliated duct, which is generally assumed to be a derivative of the gonad. Gonad lining and sperm ultrastructure are little known in heteronemerteans, a group of nemerteans belonging to the Anopla, one of the two large nemertean subgroups. Reproduction biology in heteronemertean Lineus viridis allows predicting a modified sperm type, so-called introsperm for this taxon. Nothing is known on the fate of the testes at the end of the reproductive period of this perennial species. In order to test the predictions and to broaden the data base, males of L. viridis were collected at different times of the year. Histological and ultrastructural data show that the gonad wall is lined with different aciliated endothelial cells and germ cells, while the gonoduct is formed by densely ciliated cells. The testes are completely filled with sperm cells during maturity; there is no hint at ongoing spermiogenesis at this time. The sperm consists of head, midpiece and tail. Externally, head and midpiece cannot be discriminated. The acrosome is cup-shaped and lies anterior to the nucleus which contains 6–8 lateral ridges. Three long mitochondria mark the midpiece. They line the posterior section of the nucleus and extend up to the level of the ciliary basal structures. The sperm morphology corroborates the predictions derived from the mode of reproduction. At the end of the reproductive period the male gonads change cellular composition, while the gonoduct degenerates. Provided that both sexes show the same growth rate, male offspring acquire sexual maturity earlier than female offspring, since L. viridis males are always smaller than the females. In contrast to the males, females keep their gonads and gonoducts during most time of the year. Since large males were never found within the studied population, these data indicate that L. viridis might be a consecutive hermaphrodite.     

Vergleichende Ultrastrukturuntersuchungen der Eu- und Paraspermien von 13Protodrilus-Arten (Polychaeta,Annelida) und ihre taxonomische und phylogenetische Bedeutung

The morphology of the slender, filiform spermatozoa of 13Protodrilus species of 22 different populations is investigated by light and transmission electron microscopy. All species have two types of spermatoza: fertile euspermatozoa, and paraspermatozoa, which are probably infertile and may comprise up to 20% of the total number of mature gametes. This is the first record of sperm dimorphism in polychaetes. The general construction pattern of the euspermatozoa is very complex. It shows a longish tapering acrosomal vesicle with an internal acrosomal rod, a rod-like conical nucleus, and a midpiece with numerous very complex supporting elements and two thin mitochondrial derivatives. Further, it has a ‘peribasal body’ surrounding the basal body of the axoneme, an anulus region with an ‘anchoring apparatus’ and an anulus cuff. Posteriorly, the tail region proper contains in some species 2 to 9 supporting rods. In several species the euspermatozoon shows very distinct and species-specific alternations of this ‘general pattern’ relating to e.g. size of sperm elements, structure of acrosome and nucleus, presence or absence of axial rod, and number, shape and size of supporting elements in midpiece and tail. In a number of species some sections of the euspermatozoon overlap with each other more or less strongly. The paraspermatozoon has a comparatively simple construction pattern and possesses no supporting structures in midpiece and tail region. The midpiece is very short and, in some species, entirely surrounded by its two thin and elongate mitochondrial derivatives. An axial rod is often missing or reduced; different sperm sections never overlap each other. In contrast to the euspermatozoa, the paraspermatozoa of the different species have a very similar ultrastructure. Their possible function in spermatophore transfer and histolytical opening of the female epidermis is discussed. A comparison of the different forms of euspermatozoa inProtodrilus elucidates possible plesiomorphous and apomorphous sperm traits. Very likely, the hypothetical plesiomorphous type of spermatozoa inProtodrilus has a very similar morphology to that of the paraspermatozoa, which for this reason are considered to be a sort of persisting representatives of the ancientProtodrilus sperm type. InProtodrilus, the different traits of the euspermatozoa represent excellent taxonomic characters for distinguishing species (e.g. ‘sibling species’). They can also be used well for phylogenetics within the genus, whereas the relations ofProtodrilus to other polychaete groups cannot be clarified solely on the basis of sperm characters, since in all groups the sperm structure is primarily an adaptation to a specific mode of reproduction. Generally, the value of sperm characters in phylogenetic considerations at higher taxonomic levels seems to be very limited due to the surprisingly wide range of different sperm structures within a single genus as is demonstrated in the present paper.