Ultrastructure of spermiogenesis in Plagioscion squamosissimus (Teleostei, Perciformes, Sciaenidae)

Spermiogenesis in Plagioscion squamosissimus occurs in cysts. It involves a gradual differentiation process of spermatids that is characterized mainly by chromatin compaction in the nucleus and formation of the flagellum, resulting in the spermatozoa, the smallest germ cells. At the end of spermiogenesis, the cysts open and release the newly formed spermatozoa into the lumen of the seminiferous tubules. The spermatozoa do not have an acrosome and are divided into head, midpiece, and tail or flagellum. The spermatozoa of P. squamosissimus are of perciform type with the flagellum parallel to the nucleus and the centrioles located outside the nuclear notch.     

Ultrastructure of spermatogenesis and mature spermatozoa in the flatworm Prosthiostomum siphunculus (Polycladida,Cotylea)

This is the first study investigating spermatogenesis and spermatozoan ultrastructure in the polyclad flatworm Prosthiostomum siphunculus. The testes are numerous and scattered as follicles ventrally between the digestive ramifications. Each follicle contains the different stages of sperm differentiation. Spermatocytes and spermatids derive from a spermatogonium and the spermatids remain connected by intercellular bridges. Chromatoid bodies are present in the cytoplasm of spermatogonia up to spermatids. During early spermiogenesis, a differentiation zone appears in the distal part of spermatids. A ring of microtubules extends along the entire sperm shaft just beneath the cell membrane. An intercentriolar body is present and gives rise to two axonemes, each with a 9 + “1” micro‐tubular pattern. Development of the spermatid leads to cell elongation and formation of a filiform, mature spermatozoon with two free flagella and with cortical microtubules along the sperm shaft. The flagella exit the sperm shaft at different levels, a finding common for acotyleans, but so far unique for cotylean polyclads. The Golgi complex produces numerous electron‐dense bodies of two types and of different sizes. These bodies are located around a perinuclear row of mitochondria. The elongated nucleus extends almost along the entire sperm body. The nucleus is wide in the proximal part and becomes narrow going towards the distal end. Thread‐like chromatin mixed with electron‐dense intranuclear spindle‐shaped bodies are present throughout nucleus. The general sperm ultrastructure, the presence of intranuclear bodies and a second type of cytoplasmic electron‐dense bodies may provide characters useful for phylogenetic analysis.     

Ultrastuctural and cytochemical study of spermatogenesis in Scrobicularia plana (Mollusca,Bivalvia)

The ultrastructure of the mature spermatozoa and spermatogenesis of the bivalve Scrobicularia plana are described. Support cells extend from the basal lamina to the lumen of the testis and are laterally connected to the germinal epithelium. Germ cells present intercellular bridges and flagella since the spermatogonial stage. While spermatogonia and spermatocytes appear connected to support cells by desmosome-like junctions, elongated spermatids are held at the acrosomal region by support cell finger-like processes. During spermiogenesis, the acrosomal vesicle differentiates from a golgian saccule and then migrates to the nuclear apex. A microtubular manchette arising from centrioles surrounds the acrosomal vesicle, the nucleus, and the mitochondria at the time these three organelles start their elongation, disappearing after that. The mature spermatozoon of S. plana lacks a distinct midpiece because the mitochondria extend from the region of the pericentriolar complex along the nucleus anteriorly for approximately 1.4 μm. The features of this bivalve type of modified spermatozoon are compared with those of other animal groups having similar modifications.     

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.     

Ultrastructure of spermiogenesis in Plagioscion squamosissimus (Teleostei, Perciformes, Sciaenidae)

Spermiogenesis in Plagioscion squamosissimus occurs in cysts. It involves a gradual differentiation process of spermatids that is characterized mainly by chromatin compaction in the nucleus and formation of the flagellum, resulting in the spermatozoa, the smallest germ cells. At the end of spermiogenesis, the cysts open and release the newly formed spermatozoa into the lumen of the seminiferous tubules. The spermatozoa do not have an acrosome and are divided into head, midpiece, and tail or flagellum. The spermatozoa of P. squamosissimus are of perciform type with the flagellum parallel to the nucleus and the centrioles located outside the nuclear notch.     

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.     

Ultrastructure of spermiogenesis in the gastropod Calliotropis glyptus Watson (Prosobranchia: Trochidae)with special reference to the embedded acrosome

Testicular spermatozoa and sperm development in the archaeogastropod Calliotropis glyptus Watson (Trochoidae: Trochidae) are examined using transmission electron microscopy and formalin-fixed tissues. During spermiogenesis, the acrosome, formed evidently through fusion of Golgi-derived proacrosomal vesicles, becomes deeply embedded in the condensing spermatid nucleus. Two centrioles (proximal and distal), both showing triplet microtubular substructure, are present in spermatids—the distal centriole giving rise to the sperm tail and its associated rootlet. During formation of the basal invagination in the spermatid nucleus, centrioles, and rootlet move towards the nucleus and come to lie totally within the basal invagination. Mitochondria are initially positioned near the base of the nucleus but subsequently become laterally displaced. Morphology of the mature spermatozoon is modified from that of the classic primitive or ect-aquasperm type by having 1) the acrosome embedded in the nucleus (the only known example within the Mollusca), 2) a deep basai invagination in the nucleus containing proximal and distal centrioles and an enveloping matrix (derived from the rootlet), 3) laterally displaced periaxonemal mitochondria, and 4) a tail extending from the basal invagination of the nucleus. Implantation of the acrosomal complex and centrioles within imaginations of the nucleus and lateral displacement of mitochondria effectively minimize the length of the sperm head and midpiece. Such modifications may be associated with motility demands, but this remains to be established. The unusual features of C. glyptus spermatozoa, though easily derivable from ‘typical’ trochoid sperm architecture, may prove useful in delineating the genus Calliotropis or tracing its relationship to other genera within the trochid subfamily Margaritinae.     

On sperm ultrastructure,spermiogenesis and the spermatophore of Heterolaophonte minuta (Copepoda,Harpacticoida)

Summary The spermatophore, mature spermatozoon and spermiogenesis of Heterolaophonte minuta have been investigated by light and electron microscopy. The spermatophore contains three different secretions which are responsible for the discharge of the contents of the spermatophore, the formation of the fertilization tube and the storage of the spermatozoa. The spermatozoon represents a type new for the Copepoda. It is a filiform cell about 25 m in length, ellipsoid in transverse section and tapered at the posterior end. The elongated nucleus contains chromatin fibrils and does not possess a nuclear envelope. Posterior to the nucleus, six mitochondria are placed one after the other. The posterior part of the spermatozoon contains parallel pseudomembranes. The gamete is not helically twisted and is without a flagellum and centrioles. The most remarkable feature of the spermatozoon is an osmiophilic cap in front of the nucleus. This cap corresponds to the acrosome of the spermatozoon. Early stages of spermiogenesis take place in the testis, where the spermatids are incorporated into accessory cells. The origin of the chromatin fibrils and the glycocalyx, as well as the breakdown of the nuclear envelope and centrioles, represent the final steps of spermiogenesis which occur in the vas deferens.     

Identification of TMCO2 as an acrosome‐associated protein during rat spermiogenesis

We isolated the transmembrane and coiled‐coil domains 2 (Tmco2) gene using a polymerase chain reaction‐based subtraction technique. Tmco2 is predominantly expressed in rat testes starting from 4 weeks of age. Rat TMCO2 consists of 187 amino acids with a predicted molecular mass of 20.6 kDa. When expressed in COS7 cells, TMCO2 was found as vesicle‐like structures in the cytoplasm, whereas TMCO2ΔTM lacking the transmembrane (TM) region was found diffused in the cytoplasm. These results suggest that the TM region in TMCO2 is essential for its specificity of localization. Immunocytochemical analyzes indicated that rat TMCO2 was localized as small semiluminate bodies or cap‐like structures in the vicinity of round spermatid nuclei and as curved lines associated with nuclei of elongated spermatids and caput epididymal spermatozoa. However, it was detected in only a small part of cauda epididymal spermatozoa. Double immunolabeling of the spermatids and spermatozoa with the anti‐TMCO2 antibody and the monoclonal anti‐MN7 antibody showed that TMCO2 was predominantly associated with the inner acrosomal membrane in spermatids and caput epididymal spermatozoa. Our findings suggest that TMCO2 might be involved in the process of acrosome biogenesis, especially binding of acrosome to a nucleus, during spermiogenesis.     

The aberrant spermatogenesis of the Haplothrips simplex (Buffa) (Thysanoptera): ultrastructural study

The aberrant spermatogenesis of the haploid insect Haplothrips simplex (Thysanoptera) is described. The process, which occurs in the pupal instars, is characterized by two mitotic divisions, the second of which gives rise to two different-sized spermatids: the larger spermatids have a nucleus with diffuse chromatin and proceed into spermiogenesis, while the small spermatids have pycnotic nuclei and degenerate. Both types of spermatids contain two centrioles parallely rather than orthogonally oriented. The occurrence of two centrioles supports a close relationship between Thysanoptera and Phthyraptera. Before the beginning of spermiogenesis, however, the functional spermatids show the unusual presence of a third parallel centriole which is formed by the duplication of one of the two pre-existing centrioles.     

Spermatogenesis revisited

Summary Gametogenesis is normal through meiosis and the earliest phases of spermiogenesis in male-sterile mice homozygous for the recessive, pleiotropic, mutation hpy (hydrocephalic-polydactyl). However, structurally complete sperm flagella were not encountered. Instead, partially assembled axonemal structures and/or poorly organized aggregates of other tail components (mitochondria, outer coarse fibers) were seen at the posterior poles of nuclei in older spermatids. The ultrastructure of centrioles in spermatids was normal, but that of axonemes associated with them was not. These findings suggest that the observed flagella dysgenesis results from defects in assembly rather than from defective intiation centers. Released gametes usually consisted of distorted nuclei and associated acrosome enclosed in a relatively close fitting plasma membrane. Perturbations of sperm head development were also encountered; they included extreme nuclear elongation, and distortion of the acrosome and underlying nuclear material by inpushings of finger-like processes of Sertoli cells. It is believed that sperm head anomalies are secondary consequences of the mutant condition. The findings support the view that the hpy locus represents one of a number of genes primarily involved in the mediation of flagella development.Aided by a grant from The National Foundation-March of DimesA preliminary account of some of the findings was presented elsewhere (Bryan, 1976)The author wishes to express his thanks to Mrs. Charla Danforth and Mr. James Barrows for capable technical assistance, to Dr. Walter J. Humphreys, Director, The Central Electron Microscope Laboratory, University of Georgia, for use of the facility, and to Dr. Willard F. Hollander, Iowa State University, for original breeding stock and helpful discussions     

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.     

The spermatogenesis and the sperm structure of Terebrantia (Thysanoptera, Insecta)

Spermatogenesis and the sperm structure of the terebrantian Aeolothrips intermedius Bagnall are described. Spermatogenesis consists of two mitotic divisions; the second is characterized by the loss of half of the spermatids, which have pyknotic nuclei. Early spermatids have two centrioles, but when spermiogenesis starts, a third centriole is produced. The three basal bodies give rise to three flagella; later these fuse into a single flagellum which contains three 9 + 0 axonemes. The basal bodies are surrounded by a large amount of centriole adjunct material. During spermiogenesis this material contributes to the shifting of the three axonemes towards the anterior sperm region parallel to the elongating nucleus, and it is transformed into a dense cylinder. In the mature spermatids the three axonemes amalgamate to create a bundle of 27 doublet microtubules. Near the end of spermiogenesis the dense cylinder of the centriole adjunct lies parallel to the nucleus and the axonemes. It ends where the mitochondrion appears at half-sperm length. We confirm that Terebrantia testes have a single sperm cyst; their sperm are characterized by a cylindrical nucleus, three axonemes fused into one, a small mitochondrion and a short cylindrical centriole adjunct which corresponds to the dense body described in a previous work. The acrosome is lacking. At the midpoint of the anterior half of the sperm the outline of the cross-section is bilobed, with the nucleus contained in a pocket evagination of the plasma membrane. These characters are discussed in light of a comparison between Tubulifera and Terebrantia.     

Ultrastructure of spermiogenesis in <Emphasis Type="Italic">Cristatella mucedo</Emphasis> Cuvier (Bryozoa: Phylactolaemata: Cristatellidae)

In Cristatella mucedo spermiogenesis occurs in a morula consisting of a large number of spermatids connected with a central cytophore. The mature sperm cell is filiform and consists of a head, a midpiece and a tail region, the latter two separated by a deep circular constriction. The comparatively short head contains a drop-shaped, bilaterally symmetrical and pointed nucleus capped by a minute acrosome. The single centriole is placed in a deep posterior invagination of the nucleus followed by the axoneme with the typical 9 + 2 pattern. The elongated midpiece is 0.9–1.1 μm thick and contains several helices of mitochondria surrounding the axoneme. The tail is thicker (1.3 μm) and richer in cytoplasm with many compact accumulations of an electron-dense substance lying peripherally and another less dense material wrapped around the axoneme. The course of the spermiogenesis and the fine structure of the sperm are very similar to that of Plumatella fungosa. Comparison with other species shows that the same sperm type is recognizable in four of the five families of Phylactolaemata and, provided it occurs also in the fifth family, the Stephanellidae, is a synapomorphy of the entire class.     

Semicystic spermatogenesis and biflagellate spermatozoon ultrastructure in the Nile electric catfish Malapterurus electricus (Teleostei: Siluriformes: Malapteruridae)

Spermatogenesis and spermatozoon ultrastructure in the Nile electric catfish Malapterurus electricus are described using scanning and transmission electron microscopy. Although the testis organization conforms to the ‘unrestricted’ spermatogonial type, the species has a rare type of spermatogenesis not previously described among catfishes, ‘semicystic’, in which the cyst ruptures before the spermatozoon stage. Spermiogenesis also involves some peculiar features such as condensation of the chromatin in the posterior part of the nucleus to form a compact electron‐dense mass with some irregular electron‐lucent lacunae, while the uppermost part of the nucleus is a loose electron‐lucent area, absence of the nuclear rotation and, as a consequence, the centriolar complex and the initial segment of each flagellum arise directly in a position perpendicular to the basal pole of the nucleus, and occurrence of numerous vesicles in the midpiece. In addition, spermiogenesis includes migration of the diplosome and mitochondria to the basal pole of the nucleus, formation of two moderate nuclear fossae, each of which contains the centriolar complex, development of two independent flagella and elimination of the excess cytoplasm. The mature spermatozoon has a more or less round head with no acrosome or acrosomal vesicle, a long midpiece with numerous mitochondria and vesicles and two long tails or flagella having the classical axoneme structure of 9 + 2 microtubular doublet pattern and with no lateral fins and membranous compartment. These findings suggest that the ultrastructural features of spermiogenesis and spermatozoa of M. electricus are synapomorphies of types I and II spermiogenesis and spermiogenesis is closely similar to the type described in the Nile catfish Chrysichthys auratus.     

OBSERVATIONS ON THE FINE STRUCTURE OF THE DEVELOPING SPERMATID IN THE DOMESTIC CHICKEN

The kinetic apparatus, the acrosome and associated structures, and the manchette of the spermatid of the domestic chicken have been studied with the electron microscope. The basic structural features of the two centrioles do not change during spermiogenesis, but there is a change in orientation and length. The proximal centriole is situated in a groove at the edge of the nucleus and oriented normal to the long axis of the nucleus and at right angles to the elongate distal centriole. The tail filaments appear to originate from the distal centriole. The plasma membrane is invaginated along the tail filaments. A dense structure which appears at the deep reflection of the plasma membrane is identified as the ring. The fine structure of the ring has no resemblance to that of a centriole and there is no evidence that it is derived from or related to the centrioles. The tail of the spermatid contains nine peripheral pairs and one central pair of tubular filaments. The two members of each pair of peripheral filaments differ in density and in shape: one is dense and circular, and the other is light and semilunar in cross-section. The dense filaments have processes. A manchette consisting of fine tubules appears in the cytoplasm of the older spermatid along the nucleus, neck region, and proximal segment of the tail. The acrosome is spherical in young spermatids and becomes crescentic and, finally, U-shaped as spermiogenesis proceeds. A dense granule is observed in the cytoplasm between acrosome and nucleus. This granule later becomes a dense rod which is interpreted as the perforatorium.     

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.     

Studies on the fine structure of spermatids and spermatozoa of the crab, Pinnixia sp

Early spermatids of the carb, Pinnixia sp., are characterized by a large nuclear/cytoplasmic ratio. During early spermiogenesis an organelle, composed of pentalaminar membranes derived from the endoplasmic reticulum and nuclear membranes, is formed. Portions of this organelle become incorporated within a cylinder-shaped invagination of the acrosome. Portions of the nuclear membrane disappear beneath the acrosome resulting in intermingling of nucleoplasm, centrioles and mitochondria. The nuclear membrane elsewhere is found as a pentalaminar membrane underlying the plasma membrane. Centrioles are found in mature spermatozoa at the base of the cylinder-shaped acrosomal invagination, and mitochondria are found intermingled with nuclear remanants surrounding the acrosome. This datum is compared to previously described events in spermiogenesis of other decapod crustacea and arthropods exhibiting similar modes of spermiogenesis. It is concluded that the differences and/or similarities exhibited by centrioles and mitochondria in these forms could be significant in terms of subsequent zygote differentiation.     

Behaviors of nucleus, basal bodies and microtubules during eupyrene and apyrene spermiogenesis in the silkworm, Bombyx mori (Lepidoptera)

Observations with immunostaining for tubulin and electron microscopy revealed that silkworm eupyrene spermiogenesis was characterized by an attachment of the basal body to the nucleus except in the period of movement for unidirectional arrangement. In young eupyrene sperm, a microtubule basket caught the nucleus, which thereafter was transformed elliptically. Microtubules were also observed along the elongated acrosome and mitochondrial derivatives. During apyrene spermiogenesis, however, the basal body was not attached to the nucleus and approached the head cyst cell after the completion of unidirectional arrangement, leaving the round nucleus in the middle of the cell. The presence or absence of the phenomenon in which the basal body attaches to the nucleus seems to be essential in the course of diverse spermatogenesis of the silkworm.     

STUDIES OF SPERMIOGENESIS IN A NEMATODE, NIPPOSTRONGYLUS BRASILIENSIS

The fine structure of the developing spermatids and the mature sperm of Nippostrongylus brasiliensis was investigated. Immature spermatids are found at one end of the tubelike testis, and the mature sperm at the other. The spermatid has a prominent nucleus, with the chromatin clumped at the margin. It also contains a pair of centrioles, located near the nucleus. The cytoplasm is filled with ribosomal clusters, but it lacks an organized Golgi area or endoplasmic reticulum. Besides the normal mitochondria, the spermatid has specialized mitochondrionlike inclusions with dense matrix, few broad cristae, and a crystalloid structure always facing the nucleus. As spermiogenesis proceeds, the nucleus elongates, comes to lie at one end, and later evaginates to form a separate head structure, leaving the mitochondria and other cytoplasmic organelles in a broad cytoplasmic region. The nuclear material becomes filamentous and spiral, and the centrioles come to lie at one end near the junction of the head and the cytoplasmic portion of the sperm. Microtubules are found in the cytoplasmic region extending from the tubelike nucleus. The specialized mitochondria are about eighteen in number, and are arranged in rows in staggered groups of three around the microtubules in the cytoplasmic region. The mature sperm is aflagellate and lacks an acrosome. No movement of the sperm was ever observed.