Sperm Bundles in the Seminal Vesicle of the Crematogaster victima (Smith) Adult Males (Hymenoptera: Formicidae)

This study establishes the presence of spermatodesm in the seminal vesicles of sexually mature males of Crematogaster victima (Smith). In this species, the spermatozoa are maintained together by an extracellular matrix in which the acrosomal regions are embedded. This characteristic has not yet been observed in any other Aculeata. However, the sperm morphology in this species is similar to that described for other ants. The spermatozoa measure on average 100 μm in length, and the number of sperm per bundle is up to 256. They are composed of a head formed by the acrosome and nucleus; this is followed by the flagellum, which is formed by the centriolar adjunct, an axoneme with a 9?+?9?+?2 microtubule pattern, two mitochondrial derivatives, and two accessory bodies. The acrosome is formed by the acrosomal vesicle and perforatorium. The nucleus is filled with compact chromatin with many areas of thick and non-compacted filaments. Both mitochondrial derivatives have the same shape and diameters. The presence of sperm bundles in sexually mature males differentiates C. victima from other ants; however, the similarities in the sperm ultrastructure support the monophyly of this insect group.     

Sperm structure of Mecoptera and Siphonaptera (Insecta) and the phylogenetic position of<Emphasis Type="Italic"> Boreus hyemalis</Emphasis>

Sperm ultrastructure has been studied in three species of the taxa Mecoptera and Siphonaptera. The spermatozoon of the scorpion fly Panorpa germanica shows an apical bilayered acrosome, a helicoidal nucleus, a centriolar region and a 9+2 flagellar axoneme helicoidally arranged around a long mitochondrial derivative. A second mitochondrial derivative is very short and present only in the centriolar region. A single accessory body is present and it is clearly formed as a prolongation of the centriole adjunct material. Two lateral lamellae run parallel to the nucleus. The snow fly Boreus hyemalis has a conventional sperm structure and shows a bilayered acrosome, a long nucleus, a centriolar region, two mitochondrial derivatives and two accessory bodies. The axoneme is of the 9+2 type and is flattened at the tail tip. Both P. germanica and B. hyemalis have two longitudinal extra-axonemal rods and have a glycocalyx consisting of longitudinal parallel ridges or filaments. The spermatozoon of the flea Ctenocephalides canis has a long apical bilayered acrosome, a nucleus, a centriolar region, a 9+2 axoneme wound around two unequally sized mitochondrial derivatives, and two triangular accessory bodies. In the posterior tail end the flagellar axoneme disorganises and a few microtubular doublets run helicoidally around the remnant mitochondrial derivative. The glycocalyx consists of fine transverse striations. In all three species, the posterior tail tip is characterised by a dense matrix embedding the disorganised axoneme. From this comparative analysis of the sperm structure it is concluded that Mecoptera, as traditionally defined, is monophyletic and that B. hyemalis is a member of Mecoptera rather than of Siphonaptera.     

Structure and ultrastructure of the spermatozoa of Trichogramma pretiosum Riley and Trichogramma atopovirilia Oatman and Platner (Hymenoptera: Trichogrammatidae)

Lino‐Neto, J., Báo, S. N. and Dolder, H. 2000. Structure and Ultrastructure of the Spermatozoa of Trichogramma pretiosum Riley and Trichogramma atopovirilia Oatman and Platner (Hymenoptera: Trichogrammatidae). —Acta Zoologica (Stockholm) 81 : 205–211 Spermatozoa of the Trichogramma pretiosum and T. atopovirilia are very slender and long, about 0.35 µm in diameter and 283 µm and 106 µm in length, respectively. Under light microscopy, they appear wavy along their entire length. The head contains a small acrosome which, together with the initial nuclear region is surrounded by an ‘extracellular sheath’, from which innumerable filaments irradiate. The nucleus is filled with homogeneous, compact chromatin and is attached to the flagellum by an electron dense centriolar adjunct, which extends anteriorly from the nuclear base. The flagellum consists of an axoneme with the 9 + 9 + 2 microtubule arrangement pitched in a long helix, as well as a pair of spiralling mitochondrial derivatives which coil around the axoneme. Based on these characteristics, the sperm of these Trichogramma are very similar to the chalcidoids studied to date and differ from non‐chalcidoid Hymenoptera. They differ widely from the sperm of T. dendrolimi and T. ostriniae studied, where no helically twisted structure is shown. However, based on these results we argue that the spiralling of the flagellar structures is a synapomorphy for Trichogrammatidae as well as for Eulophidae + Eurytomidae + Pteromalidae.     

Structure and ultrastructure of the spermatozoa of Bephratelloides pomorum (Fabricius) (Hymenoptera: Eurytomidae)

The spermatozoa of Bephratelloides pomorum are very long and fine. Each spermatozoon measures about 620 μm in length by 0.38 μm in diameter and, when seen under the light microscope, appears to be wavy along its entire length. The head, which is approximately 105 μm, comprises a small acrosome and a nucleus. The acrosome is made up of a cone-shaped acrosomal vesicle surrounding the perforatorium and the anterior end of the nucleus. Innumerable filaments radiate from it. The perforatorium has a diameter equal to that of the nucleus at their junction, where it fits with a concave base onto the rounded nuclear tip. The nucleus is helicoidal and completely filled with homogeneous compact chromatin. It is attached to the tail by a very long and quite electron-dense centriolar adjunct that extends anteriorly from the centriole in a spiral around the nucleus for approximately 8.5 μm. The tail consists of an axoneme with the 9+9+2 microtubule arrangement pitched in a long helix, as well as a pair of spiraling mitochondrial derivatives (with regularly arranged cristae) that coil around the axoneme, and two small accessory bodies. As well as the spiraling of the nucleus, mitochondrial derivatives and axonemal microtubules, the sperm of B. pomorum present other very different morphological features. These features include the acrosome and centriolar adjunct, both of which differentiate the spermatozoa from the majority of sperm found in other Hymenoptera. In addition these structural variations demonstrate that the sperm of chalcidoids provide characteristics that can certainly prove useful for future phylogenetic analysis at the subfamily level and, possibly, the genus too.     

Sperm morphology of the Prorops nasuta (Waterston, 1923) (Hymenoptera: Bethylidae)

In the present study, spermatozoa of the Prorops nasuta (Hymenoptera: Bethylidae) parasitoid were described morphologically. This is the first publication to describe a species belonging to the superfamily Chrysidoidea. Light and transmission electron microscopy were used. The spermatozoa of P. nasuta are linear, with a mean length of 665 μm. The acrosome is composed of an acrosomal vesicle and a perforatorium. The nucleus measures approximately 17 μm in length and is circular at its cross-section; however, its anterior extremity is oval. The chromatin is electron-dense and compact, although there are clear areas in the posterior peripheral regions. In the nucleus-flagellum transition region, the cross-section of the centriole adjunct is oval, with a pleated border and an E-PTA-positive peripheral region. The axoneme shows a 9 + 9 + 2 microtubule arrangement. The microtubules are E-PTA positive and, at the posterior extremity, the accessories are the last to terminate. The diameters and shapes of the two mitochondrial derivatives are almost identical. One begins beside the nuclear base and the other after the centriole adjunct. Posteriorly, they terminate together, immediately before the axoneme. Both have mitochondrial cristae and a region of paracrystalline material; however, the format and arrangement of this material differs from those of all other species previously studied. The paracrystalline material is more strongly E-PTA positive than the cristae region. Accessories bodies are electron-dense and located between the mitochondrial derivatives and the axoneme. In general, P. nasuta spermatozoa are similar to those of the majority of Hymenoptera; however, they have various exclusive characteristics that may be useful for studying the phylogeny and taxonomy of the superfamily Chrysidoidea and of Hymenoptera in general.     

Sperm structure and spermiogenesis in Atelura formicaria Heyden (Zygentoma, Insecta)

The spermatozoon of Atelura formicaria (Zygentoma) shows several features that are typical of insects: an apical acrosome, an elongated dense nucleus, a centriole with expanded centriolar adjunct material, two large mitochondrial derivatives, and two thin accessory bodies located beneath the nucleus. The axoneme exhibits a 9 + 9 + 2 pattern with accessory tubules formed by 16 protofilaments and intertubular material. However, spermatozoa of A. formicaria show some remarkable features. The sperm cell is short for an insect, being only 50 µm in length. The nucleus is characterized by the presence of two lateral grooves which are filled with numerous infoldings of the nuclear envelope. In a cross-section the chromatin has the configuration of the Leonardo da Vinci's 'Vitruvian man'. Each mitochondrial derivative has a peculiar structure with peripheral cristae and four crystalline bodies in its matrix; two of these crystalline bodies are large and have differently orientated cristal planes. At the end of spermiogenesis, sperm bundles are stored in the proximal part of the testes. Secretions from the epithelial wall of this region give rise to large globular structures. These include sperm bundles intermingled with dense granules, forming the so called 'spermatolophids'. These formations descend along the deferent duct and are stored in the expanded seminal vesicle. Atelura spermatozoa do not pair as in some Lepismatidae, nor do they fuse as in Tricholepidion (Lepidotrichidae). Thus, sperm aggregation in Zygentoma is realized according to different modalities and can hardly be considered as a synapomorphic trait of its subtaxa.     

Comparative ultrastructure of ant spermatozoa (Formicidae: Hymenoptera)

Mature spermatozoa from spermathecae of founding queens were obtained from 5 species of ants, representing the major subfamilies Myrmicinae (Acromyrmex versicolor, Crematogaster sp.) and Dolichoderinae (Tapinoma sessile, Conomyrma insana, Conomyrma wheeleri). The ultrastructure of ant spermatozoa has many features in common with that of higher insects and is similar to that of other Hymenoptera. Structural similarities to spermatozoa of other Hymenoptera include an acrosome containing an internal rod that extends into the nucleus, two elongate mitochondrial derivatives, a centriolar adjunct, and an axonemal arrangement of 9 + 9 + 2 that includes well-developed coarse, or accessory, tubules. Spermatozoa obtained from A. versicolor, a species that is known to store and utilize viable sperm from this supply for over 10 years, show greater development of the mitochondrial derivatives than do the other species. The most distinctive feature of ant spermatozoa in comparison to other Hymenoptera is the large size of the centriolar adjunct relative to the other organelles. The centriolar adjunct is located posterior to the nucleus, anterior to the mitochondrial derivatives, and opposite the axoneme.     

Morphology of the male reproductive system and sperm ultrastructure of Leucoptera coffeella (Lepidoptera: Lyonetiidae)

The male reproductive tract of Leucoptera coffeella was processed for light and transmission electron microscopy. In the testis, the eupyrene cells are arranged in individual cysts, while the apyrene cysts form aggregates, never observed in other Lepidoptera. Both cysts contain 128 spermatozoa, which differ from the typical pattern. In the seminal vesicle, both types of spermatozoa are dispersed in the lumen, also different from other Lepidoptera. The apyrene spermatozoa are similar to those observed for other Lepidoptera. They present an anterior region covered by a dense cap and the flagellum is composed of a 9 + 9 + 2 axoneme and two mitochondrial derivatives. The eupyrene spermatozoa, however, differ from the typical pattern for Lepidoptera. Their anterior region contains a nucleus, an acrosome and a peculiar arc of eight accessory microtubules connected to the plasma membrane by dense bridges. In the nucleus–flagellum region, the ninth accessory microtubule is assembled between both mitochondrial derivatives, to participate in the axoneme. The flagellum comprises a 9 + 9 + 2 axoneme and two mitochondrial derivatives with paracrystalline cores. External to the plasma membrane and close to the accessory microtubules, there are tufts of an amorphous material, suggesting reduced lacinate appendages, while the reticular ones are absent. The reduction of lacinate appendages and the absence of sperm bundles in the seminal vesicle support the concept that the appendages of other Lepidoptera could be associated with the eupyrene aggregations. The characters ‘number of spermatozoa per cyst’ and ‘absence of bundles’ should be considered plesiomorphic, supporting the position of this taxon in the base of the Ditrysia.     

Ultrastructural characterization of spermatozoa in euglossine bees (Hymenoptera, Apidae, Apinae)

Summary. Euglossine spermatozoa are the longest described to date for the Hymenoptera. This cell includes a head and a flagellar region. In transverse sections, the acrosome is circular at the tip but has an oval contour along most of its length. The perforatorium penetrates into a deep cavity in the nuclear tip. The flagellum consists in an axoneme, a pair of mitochondrial derivatives, a centriolar adjunct and a pair of accessory bodies. The axoneme has a 9+9+2 microtubule pattern which becomes gradually disorganized in the final portion, with the central microtubules and the nine doublets terminating simultaneously, followed by the accessory microtubules. The mitochondrial derivatives are asymmetric both in length and diameter. Sectioned transversally, the derivatives are ellipsoidal or have a pear shape. The larger one has a more obvious paracrystalline region. The centriolar adjunct begins at the nuclear base and extends parallel to the axoneme until it encounters the smaller mitochondrial derivative, on which it fits, making a concave groove. In addition to these consistent euglossine features, species-specific differences that might be useful in phylogenetic work on the group are also noted.Received 18 October 2003; revised 4 September 2004; accepted 4 October 2004.     

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.     

The short spermatodesm of Arge pagana (Hymenoptera: symphyta)

In the seminal vesicle of the 'symphyta'Arge pagana the spermatozoa are stored in motile spermatodesm bundles, maintained by an anterior cap of extracellular material. This cap consists of a denser cortex and of an internal matrix, where part of the sperm heads are embedded. The number of spermatozoa per bundle is variable. The spermatozoa are short, only 30microm long, with a head region of about 23microm, and a very short flagellum of about 7microm. The head includes the acrosome, with a perforatorium, and the nucleus. The flagellum consists of an axoneme, with a 9+9+2 microtubule pattern, a centriolar adjunct, two mitochondrial derivatives and two accessory bodies. The mitochondrial derivatives are very slender and of different lengths. The longer begins at the base of the nucleus, while the shorter one starts just below the base of the centriolar adjunct. This latter is asymmetric and appears at the nuclear base, extending parallel to the axoneme up to the anterior end of the smaller mitochondrial derivative. The short spermatodesmata and the small mitochondrial derivatives characterize the A. pagana sperm. In addition, the centriolar adjunct asymmetry and the occurrence of spermatodesm bundles might be considered plesiomorphic states present in the basal Tenthredinoidea.     

Polymorphism of spermatozoa in Largus rufipennis Laporte 1832 (Heteroptera: Pyrrhocoroidea: Largidae)

The production of polymorphic spermatozoa has been registered in various insect orders such as Diptera, Lepidoptera, and Hemiptera. In this work, morphology of two types of spermatozoa produced by Largus rufipennis was reported for the first time in the Largidae family. For this, techniques including optical and transmission electron microscopy were used. Spermatozoa measured, on the average, 260 and 200 μm, and both types possessed a nucleus measuring on the average 65 μm. No ultrastructural differences were observed between the two spermatozoa types from L. rufipennis. The head region is composed of an acrosome, a nucleus, and part of the centriolar adjunct. The centriolar adjunct is in parallel with the nucleus and followed by mitochondrial derivates. The flagellum consists of an axoneme (9 + 9 + 2 microtubules) and two mitochondrial derivatives; no other accessory bodies were observed. The mitochondrial derivatives are symmetric in size and diameter. A similar quantity of the two spermatozoa types was observed in the seminal vesicle (57% of the large type and 43% of the small type), while in the spermatheca of the female, the larger spermatozoa were preferentially stored (87%). These results permit discussions concerning of the species biology reproduction, most specifically sperm competition strategies.     

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).     

Comparative spermatology of four species of strepsiptera and comparison with a species of primitive coleoptera (Rhipiphoridae)

The comparative spermatology of 4 strepsipteran species, namely, Xenos moutoni De Buysson, Elenchus tenuicornis (Kirby), Elenchus japonicus Esaki and Hashimoto and Halictophagus chilensis Hofmann, have been studied by transmission electron microscopy. The spermatozoon of all 4 species examined were seen to have: an elongated head characterized by a nucleus with an internal channel of uncondensed chromatin, a one-layered acrosome, and a tail containing a 9 + 9 + 2 axoneme and 2 partially crystallized mitochondrial derivatives. Each sperm, nevertheless, shows a few peculiarities that confirm the taxonomic value of comparative spermatology. In addition, the strepsipteran spermatozoa were seen to have a different organization to that of Pelecotoma fennica Pewkull (Rhipiphoridae, Pelecotominae), which has the typical structure of a coleopteran sperm. It was characterized by a 3-layered acrosomal complex, a tail with a 9 + 9 + 2 axoneme flanked by 2 accessory bodies and 2 mitochondrial derivatives. From the spermatological point of view, primitive Coleoptera cannot be considered to be closely related to Strepsiptera.