COMPARATIVE SPERM MORPHOLOGY OF THE PULMONATE LIMPETS TRIMUSCULUS COSTATUS, T. RETICULATUS (TRIMUSCULIDAE) AND BURNUPIA STENOCHORIAS AND ANCYLUS FLUVIATILIS (ANCYLIDAE)

Sperm ultrastructure is described for the first time in representativesof the pulmonate ‘limpet’ families Trimusculidae(Trimusculus costatus, T. reticulatus: marine) and Ancylidae(Burnupia stenochorias, Ancylus fluviatilis: freshwater). Allshow characteristic heterobranch sperm features (a spheroidalacrosomal vesicle supported by an acrosomal pedestal; a helicallykeeled nucleus and a complex, very elongate midpiece featuringparacrystalline and matrix layers sheathing the axoneme, coarsefibers and one or more glycogen helices). Posterior to the midpiece,a glycogen piece (axoneme sheathed by glycogen granules) andannulus are also present in all species. Taxonomically usefuldifferences in the shape and dimensions of the acrosome, nucleusand midpiece occur between the species. Results support thedecision of recent workers to transfer the Trimusculidae fromthe Siphonarioidea to a separate superfamily Trimusculoidea(characteristic sperm features including: narrow acrosomal pedestaloverlapping with nuclear apex; heavily keeled nucleus; midpiecewith strongly projecting secondary and glycogen helices). Therelationship of the Trimusculoidea to other pulmonates, as indicatedby sperm ultrastructure, remains uncertain largely because comparativedata for several important groups are unavailable. Spermatozoaof the two ancylids most closely resemble those of other investigatedplanor-boideans and to a lesser extent, those of the Lym-naeoidea.However, differences between Burnupia stenochorias (unique(?)accessory structure on the acrosomal pedestal; glycogen wedgeswithin the nuclear fossa; other features similar to planorbids)and Ancylus fluviatilis (all sperm features very similar toplanorbids) suggests that these patelliform ancylids are notclosely related. (Received 20 November 1997; accepted 23 January 1998)     

TRANSFER OF THE GASTROPOD FAMILY PLESIOTROCHIDAE TO THE CAMPANILOIDEA BASED ON SPERM ULTRASTRUCTURAL EVIDENCE

Using sperm ultrastructure the systematic placement and affinitiesof the caenogastropod family Plesiotrochidae are re-examined.The simultaneous hermaphrodite, Plesiotrochus crinitus Thiele,1930, produces both euspermatozoa (uniflagellate, fertile sperm)and paraspermatozoa (bi- or triflagellate, infertile sperm).Features of each type of sperm clearly indicate that the Plesiotrochidaeare closely related to the Campanilidae (Campaniloidea) andare not, as previously believed, referable to the superfamilyCerithioidea. Significant sperm synapomorphies of Plesiotrochus(Plesiotrochidae) and Campanile (Campanilidae) include the morphologyof the eusperm midpiece (seven to nine straight mitochondriasurrounded by a segmented, accessory sheath of membrane-boundvesicles) and morphology of the anucleate parasperm head (axialcore of mitochondria surrounded by a bilaterally symmetricalarrangement of axonemes and dense vesicles). The characteristicsubstructure of the cerithioidean eusperm midpiece (four straightmitochondria each containing parallel, cristal plates) is notobserved in Plesiotrochus or Campanile. Euspermatozoa of Plesiotrochusdiffer from Campanile principally in details of the acrosomalcomplex (Plesiotrochus with apical bleb, probable absence ofan accessory membrane; Campanile without apical bleb, accessorymembrane well developed), the transverse profile of all midpiecemitochondria (thin in Plesiotrochus; thick in Campanile), andmorphology of the annulus (double ring in Plesiotrochus; singlering in Campanile). In addition, all observed paraspermatozoaof Plesiotrochus are anucleate, whereas in Campanile anucleateand nucleate paraspermatozoa are present. On the basis of spermsynapomorphies of Plesiotrochus and Campanile, the Plesiotrochidaeare transferred from the Cerithioidea to the Campaniloidea. (Received 3 August 1992; accepted 18 September 1992)     

ULTRASTRUCTURE OF THE MATURE SPERMATOZOA OF THE LAND SNAIL EPIPHRAGMOPHORA TUCUMANENSIS (DOERING, 1874) (GASTROPODA: HELICOIDEA)

The ultrastructure of the sperm of Epiphragmophora tucumanensis(Doering), a pulmonate land snail belonging to the family Xanthonychidae,was examined. The results showed that this spermatozoon presents a similarmorphology to the other advanced stylommatophoran sperm described.The most peculiar characteristic is the form of the nucleus,which is straight instead of helically coiled. An acrosome ispresent, composed of an apical vesicle and an acrosomal pedestal.The connecting piece or neck region is formed by a basal bodyplus a centriolar derivative. It is also the point of originof coarse fibres that accompany the axoneme. Thus, the axonemalcomplex exhibits a 9+9+2 pattern. The midpiece is an elongatedregion composed of paracrystalline and matrix materials (collectively,the mitochondrial derivative) enclosing a single glycogen helixand the axoneme. The mitochondrial derivative extends to theposterior tip of the spermatozoon. As observed in other stylommatophorans,both an annulus and a glycogen piece are absent. (Received 25 October 1993; accepted 4 February 1994)     

SPERMOGENESIS AND SPERM STRUCTURE IN THREE SPECIES OF PATELLOIDA AND ONE SPECIES OF NIPPONACMAEA (PATELLOGASTROPODA: ACMAEOIDEA)

The spermatozoa of Patelloida profunda albonotata, P. saccharina,P. pygmaea and Nipponacmaea schrenkii (Lottiidae) are describedby transmission electron microscopy. All have ect-aquasperm,typical of invertebrates using external fertilization. The spermof all four species have a cylindrical nucleus (length: breadth> 4: 1 in P. p. albonotata and P. saccharina; <4: 1 inP. pygmaea and N. schrenkii) which tapers towards the roundedanterior end. All have an acrosome with a posterior acrosomallobe which extends into the centre of the subacrosomal space.In P. p. albonotata and P. saccharina the acrosomal contentsare undifferentiated and the posterior lobe extends to the nucleus,being separated from it by flocculent material. In P. pygmaeathe acrosomal contents are differentiated, the lobe is relativelyshort and the subacrosomal space is filled with material witha fibrous appearance. The acrosome of N. schrenkii is undifferentiatedand the posterior lobe is no more than a bulge. The sperm ofP. p. albonotata and N. sacchrina have a small (0.25 µmlong) cytoplasmic collar which surrounds the axoneme anteriorlywhereas in P. pygmaea and N. schrenkii the cytoplasmic collaris longer (1 µm) and is swollen by an electron-dense vesicle.The composition and function of this vesicle is unknown. Thespermatozoa of Patelloida and Nipponacmaea have structural featureswhich are similar to sperm of the Lottiidae providing some supportfor the placement of these genera in the Lottiidae as proposedby Lindberg & Hedegaard (1996) and Sasaki & Okutani(1993) respectively. The similarities of the sperm of P. pygmaeato N.schrenkii raise some doubts about the tax-onomic statusof the former species. Spermiogenesis in all four species issimilar to that described for other Acmaeoidea and Patelloidea.In P. pygmaea and N. schrenkii, however, in addition to theacrosomal vesicle, the Golgi body produces a number of electron-densevesicles which fuse and eventually form a single vesicle inthe collar of the mid-piece. (Received 24 October 1996; accepted 10 February 1997)     

SPERMATOZA AND SPERMIOGENESIS OF CORNIROSTRA, VALVATA AND ORBITESTELLA (GASTROPODA: HETEROBRANCHIA) WITG A DISCUSSION OF VALVATOIDEAN SPERM MORPHOLOGY

Transmission electron microscopy reveals that spermatozoa ofthe marine gastropods Cornirostra peleucida (Laseron) (Cornirostridate—genusand family newly proposed by Ponder, 1990a) and Orbitestellasp. (Orbitestellidae) closely resemble those of the freshwaterfamily valvatidae. In particular, spermatozoa of Cornirostra,Orbitestella and Valvata share a unique arrangement of helicalridges or channels within the glycogen piece. The sperm midpieceof Orbitestella and Valvata has a lamellar substructure, whilein Cornirostra thus region exhibits a single, continuous mitochondrialsheath around the axoneme. An acrosome could not be positivelyidentified in either Valvata or Orbitestella but appears tobe present in Cornirostra in the form of a hollow, irregularvesicle. Mature sperm nuclei of the three taxa are long, filiformand almost totally penetrated by the axoneme. Nuclei of Orbitestellapossess one or more helical keels (keels absent in Cornirostraand Valvata). The pattern of spermiogenesis observed suportsthe view expressed by recent authors that the Valvatoidea areclosely related to heterobrach gastropods, and should be formallytransferred to the Heterobranchia. (Received 9 September 1989; accepted 30 December 1989)     

Ultrastructure of spermiogenesis of Philippia (Psilaxis) oxytropis,with special reference to the taxonomic position of the architectonicidae (Gastropoda)

Summary Spermiogenesis of the architectonicid Philippia (Psilaxis) oxytropis was studied using transmission electron microscopy. Both spermatids and mature sperm of Philippia show features comparable to sperm/spermatids of euthyneuran gastropods (opisthobranchs, pulmonates) and not mesogastropods (with which the Architectonicidae are commonly grouped). These features include: (1) Accumulation of dense material on the outer membrane of anterior of the early spermatid nucleus — this material probably incorporated into the acrosome; (2) Structure of the unattached and attached spermatid acrosome (apical vesicle, acrosomal pedestal) accompanied by curved (transient) support structures; (3) Formation of the midpiece by individual mitochondrial wrapping around the axonemal complex, and the subsequent fusion and metamorphosis of the mitochondria to form the midpiece; (4) Presence of periodically banded coarse fibres surrounding the axonemal doublets and intra-axonemal rows of granules. A glycogen piece occurs posterior to the midpiece but is a feature observed in both euspermatozoa of mesogastropods (and neogastropods) and in sperm of some euthyneurans.Despite the lack of paracrystalline material or glycogen helices within the midpiece (both usually associated with sperm of euthyneurans), the features of spermiogenesis and sperm listed indicate that the Architectonicidae may be more appropriately referable to the Euthyneura than the Prosobranchia.Abbreviations a acrosome - ap anterior region of acrosomal pedestal - as support structures of spermatid acrosome - av apical vesicle of acrosome (acrosomal vesicle of un-attached acrosome) - ax axoneme - b basal region of acrosomal pedestal - c centriole - cf coarse fibres - cr cristal derivative of midpiece - db intra-axonemal dense granules - drs dense ring structure - gg glycogen granules - gp glycogen piece - G Golgi complex - m mitochondrion - mt microtubules - n nucleus - pm plasma membrane - sGv small Golgi vesicles     

Comparative sperm ultrastructure and spermiogenesis in basal heterobranch gastropods (Valvatoidea, Architectonicoidea, Rissoelloidea, Omalogyroidea, Pyramidelloidea) (Mollusca)

Three distinct groups of basal heterobranch gastropods (='Allogastropoda/Heterostropha') can be distinguished on the basis o f sperm and spermiogenic features: (1) Valvatoidea; (2) Architectonicoidea; (3) Rissoelloidea + Omalogyroidea + Pyramidelloidea. Sperm of pentaganglionate heterobranchs (Opisthobranchia sensu stricto + Pulmonata) conform in all respects to the pattern shown by group (3). Heterobranchia are united, and differentiated from other gastropods, by a shared suite of sperm and spermiogenic features (rounded acrosomal vesicle, nucleus usually helical, highly modified mitochondrial derivative, spermatid acrosome associated with a dense nuclear plaque, formation of the mitochondrial derivative through fusion of numerous small mitochondria along the length of the axoneme). Sperm autapomorphies of the Architectonicoidea and the Valvatoidea suggest that these two superfamilies arc distinct but terminal offshoots from ancestral heterobranchs.     

AN ULTRASTRUCTURAL STUDY OF SPERMATOZOA OF HELIX ASPERSA AND HELIX POMATIA (GASTROPODA, PULMONATA)

Spermatozoa of the pulmonates Helix aspersa Müller andH. pomatia Linnaeus are examined in detail using transmissionelectron microscopy (TEM). Important features such as the acrosome,perinuclear sheath, nucleus and terminal region of the midpieceare described for the first time. Also presented are the firstultrastructural observations on spermatozoa from spermatophoresin any pulmonate gastropod (H. aspersa). No morphological differencescould be found between sperm taken from spermatophores and thosewithin the hermaphrodite duct in H. aspersa. Spermatozoa ofH. aspersa and H. pomatia snow all the characteristics of euthyneuranspermatozoa, namely: a helically-keeled nucleus; distinctivearrangement of acrosomal components (apical vesicle, acrosomalpedestal), and extremely elongate midpiece (axoneme and glycogenhelix enclosed by matrix and paracrystalline layers). The spermnucleus of both species is short, and the midpiece also formsthe terminal portion of the spermatozoon (glycogen piece absent).The extraordinary positioning of the acrosome in H. aspersa—reflectedbackwards from the nuclear apex—is not observed in H.pomatia, though a perinuclear sheath (possibly another acrosomalcomponent) is present in sperm of both species. Helix spermatozoaare compared with other euthyneuran sperm and briefly discussedfrom the systematic viewpoint. Present address: Department of Zoology, St. Lucia, 4067, Brisbane,OLD, Australia (Received 23 May 1988; accepted 17 August 1988)     

ULTRASTRUCTURE OF SWARMERS IN THE LAMINARIALES (PHAEOPHYCEAE). II. SPERM1

The ultrastructure of sperm from 13 species in 11 genera of Laminariales collected in the northeast Pacific Ocean is unique in the brown algae. The sperm are elongate, and possess a nucleus, several mitochondria and two or three chloroplasts, but no eyespot. The anterior flagellum bears mastigonemes on the proximal half of its length; a distal “whiplash” portion lacks mastigonemes and is an extension of only the two central singlet microtubules of the axoneme. A peculiar feature of these sperm is the posterior flagellum, which is longer than the anterior flagellum and tapers distally as the doublet microtubules become singlets and decrease in number. This feature contrasts with the laminarialean zoospore, which possesses a short posterior flagellum with the usual “9 + 2” axoneme. The structure of these sperm differs from that reported for Chorda, the sperm of which resembles a primitive brown algal zoospore. The facts support the concept that Chorda is the most primitive member of the Laminariales.     

Ultrastructure of Euspermiogenesis in the Mesogastropod Stenothyra sp. (Prosobranchia, Rissoacea, Stenothyridae)

The structure of mature and developing euspermatozoa of the rissoacean gastropod Stenothyra sp. has been studied using transmission electron microscopy. During cuspermiogenesis nuclei pass through fibrillar and lamellar phases of condensation. A Golgi-derived acrosome attaches to the nucleus during the fibrillar phase. Spherical mitochondria of early euspermatids fuse to form the mitochondrial sheath which undergoes metamorphosis to form helical midpiece elements, paracrystalline material and helical midpiece compartments. Mature euspermatozoa consist of a flat acrosome (acrosomal cone, axial rod, basal plate), short curved nucleus (2.5–2.8 μm) and elongate midpiece and glycogen piece. Coarse fibres associated with the axoneme emerge from a posterior invagination of the nucleus and continue into the initial portion of the midpiece. In the proximal portion of the midpiece, two helical compartments (filled with membranous material) are present—only one of which persists further posteriorly. No compartments occur in the distal region of the midpiece. Posterior to the midpiece, the axoneme is surrounded by tightly-packed (glycogen) granules and terminates within this region. The distal end of the euspermatozoon consists solely of glycogen granules surrounded by the plasma membrane. Although coarse fibres (associated with the axoneme), midpiece paracrystalline material and helical compartments are commonly reported in sperm of euthyneuran gastropods, this represents the first report of all three features in any prosobranch euspermatozoon.     

An ultrastructural study of the spermatozoa ofEulalia sp. (Phyllodocidae),Lepidonotus sp. (Polynoidae),Lumbrineris sp. (Lumbrineridae) andOwenia fusiformis (Oweniidae)

The ultrastructure of the mature spermatozoa of four polychaetes is described:Eulalia sp. (Phyllodocidae),Lepidonotus sp. (Polynoidae),Lumbrineris sp. (Lumbrineridae) andOwenia fusiformis (Oweniidae). All the sperm show features typical of externally fertilizing sperm in having a rounded nucleus, a short unmodified midpiece, and a simple flagellum with a 9+2 axoneme.Owenia fusiformis andLepidonotus sp. have a nuclear cone extending into the subacrosomal space that may act to present the inner acrosomal membrane to the egg during fertilization. The acrosome ofLumbrineris sp. is flattened and crenulated. The sperm ofEulalia sp. is unusual in having the four mitochondria of the midpiece ensheathed by a membrane. Comparisons are made with other polychaete sperm, and the use of sperm ultrastructure as a taxonomic tool within the Polychaeta is discussed.     

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.     

SPERM COMPETITION AND THE EVOLUTION OF NONFERTILIZING SPERM IN MAMMALS

Nonfertilizing sperm with special morphologies have long been known to exist in invertebrates. Until recently, abnormal sperm in mammals were considered errors in production. Now, however, Baker and Bellis (1988, 1989) have proposed that mammalian sperm, like some invertebrate sperm, are polymorphic and adapted to a variety of nonfertilizing roles in sperm competition, including prevention of passage of sperm inseminated by another male. More specifically, their “kamikaze” sperm hypothesis proposes that deformed mammalian sperm are adapted to facilitate the formation and functioning of copulatory plugs (Baker and Bellis, 1988). Here I argue that most, maybe all, mammals are unlikely to produce nonfertilizing sperm. First, mammals might not be able to afford to evolve nonfertilizing sperm, given that a) fertilization is often unlikely despite the huge numbers of sperm produced; b) production of larger numbers of sperm is constrained, presumably because of metabolic costs, evidence for which includes the fact that in species in which sperm morphology and anatomy of the female reproductive tract increase the probability of fertilization, the numbers of sperm produced is lower than in others; and c) selection appears to act against the production of deformed sperm. Second, some of the evidence advanced for the existence of nonfertilizing sperm does not in fact support the idea. Third, accessory gland secretions are sufficient on their own to coagulate semen and produce fully functioning plugs; thus the male that used accessory gland secretions would be at a clear advantage over the male that diluted his fertilizing sperm with “kamikaze” sperm; and indeed, current evidence indicates selection on accessory glands, not sperm morphology, to enhance coagulation of semen. Fourth, predictions made on the basis of the “kamikaze” sperm hypothesis are not supported by quantitative comparisons of data from polyandrous and monandrous primates (i.e., those in which several males mate with a fertile female, and therefore in which sperm competition should be operating, and those in which only one male mates). Although sperm competition is almost certainly more intense in polyandrous genera than in monandrous genera (as indicated by, e.g., more frequent copulations and the production of more sperm per ejaculate from larger spermatogenic organs), polyandrous genera do not produce a greater proportion of deformed (i.e., nonfertilizing) sperm than do monandrous genera, or even necessarily a greater number of deformed sperm; nor a greater variety of sperm sizes—indeed they might produce fewer; nor fewer motile sperm (as might be expected if sperm are selected to stay behind and compete with sperm from subsequent males); and nor larger sperm (as might be expected if sperm are produced for functions other than to reach the egg). In sum, currently available evidence suggests that the function of all mammalian sperm is to fertilize, and that sperm competition in mammals occurs through scramble competition, not contest competition.     

SPERM MORPHOLOGY IN SERPULORBIS AND DENDROPOMA AND ITS RELEVANCE TO THE SYSTEMATIC POSITION OF THE VERMETIDAE (GASTROPODA)

Electron microscopic observations on the euspermatozoa (fertilizingsperm) and paraspermatozoa (infertile sperm) of Serpulorbissp. and Dendropoma sp. suggest that the family Vennetidae (sensustricto) is not closely allied to any true cerithioidean family.Vermetid euspermatozoa strongly resemble those of a number ofother mesogastropod superfamilies including the Stromboidea,Epitonioidea, Naticoidea, Calyptraeoidea, Cypraeoidea, somerissooideans and littorinoideans, and even some neogastropodtaxa. Vermetid paraspermatozoa are anudeate, with one or moretails extending from both ends of a central body region. Incontrast, paraspermatozoa of true cerithioideans: (1) have tailsextending from only the posterior region of the ‘head’(2) usually retain a condensed nuclear remnant and (3) sometimespossess an acrosome-like structure at the head apex. These results,in conjunction with anatomical work by Morton, suggest thatthe Vennetidae should be removed from the Cerithioidea and placedprobably in its own superfamily (Vermetoidea Raphinesque, 1815)as practised by some recent workers. The Vermetoidea possiblyare associable with the Stromboidea/Calyptraeoidea/Hipponicoidea/Xeno-phoroideablock within the Mesogastropoda ^*Present address: Department of Zoology, University of Queensland,St. Lucia, 4067, Queensland, Australia     

SPERM ULTRASTRUCTURE IN THE AUSTRALIAN OYSTER SACCOSTREA COMMERCIALIS (IREDALE & ROUGHLEY) (BIVALVIA: OSTREOIDEA)

Sperm ultrastructurc is investigated in the economically importantAustralian oyster Saccostrea commercialis (Iredale & Roughley).Mature spermatozoa consist of a broad, cap-shaped acrosomalvesicle, sub-acrosomal material (axial rod embedded in a granularmatrix), a round nucleus (deeply invaginated anteriorly), twotriplet substructure centrioles surrounded by four sphericalmitochondria, and a flagellum anchored via satellite fibresto the distal centriole and plasma membrane. Spermatozoa ofS. commercialis closely resemble those of other investigatedostreids, with the exception that the anterior region of theacrosomal vesicle is transversely banded. Differences in acrosomalsubstructure correlate with the known inability of Saccostreaand Crassostrea gametes to interact. Potential exists for useof sperm morphology in the resolution of taxonomic problemswithin the Ostreoidea. (Received 23 April 1990; accepted 28 June 1990)     

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.     

SPERM COMPETITION AND REPRODUCTIVE MODE INFLUENCE SPERM DIMENSIONS AND STRUCTURE AMONG SNAKES

The role of sperm competition in increasing sperm length is a controversial issue, because findings from different taxa seem contradictory. We present a comparative study of 25 species of snakes with different levels of sperm competition to test whether it influences the size and structure of different sperm components. We show that, as levels of sperm competition increase, so does sperm length, and that this elongation is largely explained by increases in midpiece length. In snakes, the midpiece is comparatively large and it contains structures, which in other taxa are present in the rest of the flagellum, suggesting that it may integrate some of its functions. Thus, increases in sperm midpiece size would result in more energy as well as greater propulsion force. Sperm competition also increases the area occupied by the fibrous sheath and outer dense fibers within the sperm midpiece, revealing for the first time an effect upon structural elements within the sperm. Finally, differences in male–male encounter rates between oviparous and viviparous species seem to lead to differences in levels of sperm competition. We conclude that the influence of sperm competition upon different sperm components varies between taxa, because their structure and function is different.     

An ultrastructural examination of developing and mature euspermatozoa in pyrazus ebeninus (Mollusca,Gastropoda, Potamididae)

Summary Mature and developing euspermatozoa of the prosobranch gastropod Pyrazus ebeninus, have been examined using transmission electron microscopy and phase-contrast light microscopy. The head of the mature euspermatozoon consists of a conical acrosome capping a short, rod-shaped nucleus (laterally compressed posteriorly). A basal invagination in the nucleus contains the proximal portion of the axoneme and a dense attachment matrix. Four apparently non-helical mitochondrial elements (two large, two small) comprise the midpiece each being composed of curved, inclined cristal plates and a granular matrix. The structure and arrangement of the mitochondrial elements is thus distinguishable from the helical midpiece elements found in euspermatozoa of neogastropods and most mesogastropods and possibly is widespread in the Cerithiacea. A dense ring-like structure is found closely applied to the inside of the plasma membrane at the junction of midpiece and glycogen piece.Acrosome and midpiece formation and nuclear condensation have been studied in developing euspermatozoa. Acrosome development is divided into two phases: (1) a pre-attachment phase — during which a complex early acrosome is formed often at great distance from the nuclear apex, and (2) an attachment/post-attachment phase — during which the completed preattachment phase acrosome tilts into position at the nuclear apex and subsequently elongates. The nucleus passes through a recognizable sequence of condensation phases (reticular, fibrillar and lamellar phases). Microtubules surround both the nucleus and midpiece in the final phase of maturation. The four, elongate midpiece elements of the mature euspermatozoon are apparently derived from the four large, spherical mitochondria of the euspermatid.The potential usefulness of spermatozoal ultrastructure with regard to indicating affinities between groups of gastropod families is briefly discussed.Abbreviations a acrosome - ac euspermatozoon acrosomal cone - ar euspermatozoon axial rod - ax axoneme - bp basal plate - cy cytoplasmic droplet - cs cylindrical support structures of developing acrosome - dg dense granule of pre-attachment phase developing acrosome - dp dense plates of developing acrosomal cone - g glycogen granules - gp glycogen piece - G Golgi complex - j junction of midpiece and glycogen piece - l large midpiece element - m mitochondrion - M midpiece - mt microtubules - n nucleus - pm plasma membrane - sGv small Golgi vesicles - s small midpiece element     

SPERM AND SPERMIOGENESIS IN OPISTHOTEUTHIS PERSEPHONE (OCTOPODA: CIRRATA): ULTRASTRUCTURE, COMPARISON WITH OTHER CEPHALOPODS AND EVOLUTIONARY SIGNIFICANCE

Ultrastructural features of spermiogenesis and mature spermatozoaare described for the first time in a cirrate octopod. Comparisonwith other investigated cephalopod species shows that spermatozoaof Opisthoteuthis persephone Berry most closely resemble thoseof the genus Octopus. The shared features include: (1) an elongate,solid acrosome with internal banding and a prominent helicalkeel; (2) a straight, rod-like nucleus; (3) a short midpieceand (4) a post-mitochondrial (‘annular’) skirt.Of these, the acrosomal morphology is most significant taxo-nomically,as the internal banding of the acrosomal vesicle only occursin the Octopoda. Spermatozoa of Opisthoteuthis and Octopus differhowever in the extent of the internal banding of the acrosome(poorly developed in Opisthoteuthis) and substructure of theacrosomal keel (more complex in Opisthoteuthis). Surprisingly,the extensive fibrous plug of Octopus and Vampyroteuthis sperm,is not developed either in Opisthoteuthis or in Eledone. Resultsprovide additional support for the monophyly of the Octopodabut also hint at the possibility that some groups of in-cirratesmay not be as advanced as generally believed. (Received 6 July 1992; accepted 18 August 1992)