Paraspermatogenesis of Cerithioidean snails: retention of an acrosome and nuclear remnant

Paraspermatozoa of Cerithioidea are oligopyrenic with a central nuclear remnant surrounded by glycoprotein bodies and an anterior acrosome complex. Posteriorly, the parasperm have one long and several small flagella. Biosynthesis of proteins begins in the rough endoplasmic reticulum (RER), early in paraspermatogenesis, prior to the degradation of the nucleus. The breakdown of the nucleus has features characteristic of apoptosis but a nuclear remnant remains that is composed of intact DNA. The acrosome complex of parasperm is Golgi-derived, forming posteriorly and migrating anteriorly along the plasma membrane to the apex of the nuclear remnant as the paraspermatid matures. This mechanism of acrosome formation is similar to that in euspermiogenesis in neomenioid aplacophorans and neritid snails and is plesiomorphic to mollusks. However, eusperm acrosomes of Cerithioidea form differently. In the paraspermatid, small, dense granules secreted by the Golgi body fuse to form larger granular vesicles, many of which are released by exocytosis into the seminal fluid. These granular vesicles stain red with acridine orange at pH 7 under 489-nm light, indicating that they are lysosomes. The retention of a nuclear remnant and development of an acrosome complex in the parasperm of cerithioideans, as well as the secretion of lysosomes into the seminal fluid, suggests novel functions for these unusual germ cells.     

Paraspermatogenesis in Littoraria (Palustorina) articulata, with reference to other Littorinidae (Littorinoidea, Caenogastropoda)

Abstract. The ultrastructure of paraspermatogenesis is examined in the littorinid subfamily Littorininae, with special emphasis on Littoraria (Palustorina) articulata (PHILIPPI 1846). In particular the study focuses on the fate of the nucleus and origin of the rod bodies during parasperm development. Parasperm of the Littorininae are rounded or oblong cells, which undergo an abortive meiosis and eliminate part of the nucleus but often retain a nuclear remnant. The cytoplasm is filled with numerous spherical vesicles in all Littorininae, but in Littoraria (and in certain species of Nodilittorina, Tectarius and Cenchritis) dense 'rod-bodies' also occur. Littoraria (Palustorina) are unique in possessing a flagellum-like structure termed the 'pseudotrich", which lacks an axoneme but contains microtubules during its development. Paraspermatogonia differ from euspermatogonia in the structure of the nucleus and in the extensive rough endoplasmic reticulum (RER) and swollen cytoplasm. Two types of secretions develop in Littoraria : (1) numerous, spherical granules (composed of putative glycoprotein, also seen in other Littorininae) and (2) rhomboid granules (composition uncertain but reacting positively to RNA stains; these granules arising within RER cisternae close to the nucleus). As the rhomboid granules fuse to form the larger, rod-bodies (polygonal in cross section), the RER membrane enclosing the rod-bodies becomes confluent with the outer nuclear membrane, thereby forming a common compartment. Results of this study clearly show that the rod-bodies are secretions of the RER cisternae and not, as claimed in some light microscopic accounts, the product of fusion of eusperm nuclei which have entered the parasperm cytoplasm (either by active eusperm penetration or by phagocytosis). Developmental characteristics of littorinid parasperm show differences between species and may, in some cases, provide characters diagnostic of subgenera.     

Sex and a snail's sperm: on the transport,storage and fate of dimorphic sperm in Littorinidae

Summary

Littorinid parasperm develop from a distinctive lineage of germ cells which exhibit a process of nuclear destruction that has apototic characteristics. Fragments of DNA and other nuclear breakdown products are incorporated into secretion granules in parasperm that ultimately find their way to the female bursa copulatrix. Spermatozeugmata are stored in the seminal vesicles and, if unused during the breeding season, they are recycled by phagocytosis. Attachment between eusperm and parasperm is facilitated by an electrostatic interaction of proteins. Detachment, caused by alkaline prostate fluid, occurs by the time the ejaculate reaches the tip of the penis. Thus transport of eusperm by parasperm to the female is unlikely. parasperm are sterile cells that may function in defense against rival eusperm as suggested by the presence of lysosomes, or they may act as nuptial gifts as they are packed with glycoprotein nutrients. Differences in the reactivity of different parasperm to specific lectins may enable separation of dimorphic sperm by lectin affinity chromatography, thereby facilitating future studies on individual parasperm. In female Littorinidae, sperm are stored incapacitated in storage organs, or rarely in the ovary itself. In L. littorea serotonin caused spawning of unencapsulated eggs, which, in the presence of activated sperm, became polyspermic.     

Parasperm: morphological and functional studies on nonfertile sperm

Sperm polymorphism, a phenomenon in which more than one type of sperm is produced within a species, occurs widely in animals from invertebrates to vertebrates. Sperm in this phenomenon can be categorized into fertile sperm and nonfertile sperm on the basis of fertilization ability. Nonfertile sperm can be further classified into parasperm and aberrant deformed sperm. Parasperm are sperm produced through a constant developmental process along with normal fertile eusperm, and they are readily distinguished from deformed sperm, which are irregularly crippled by unpredictable errors at certain stages during spermatogenesis. Sperm identified as parasperm occur widely in invertebrates but are presently quite limited in vertebrates. This may be the result of the deficiency both of studies on parasperm and of clear criteria to identify parasperm in vertebrates. Some vertebrates show unique spermatogenesis, such as symplastic spermatid and semicystic spermatogenesis. Thus, parasperm must be identified by comparing cells in each cyst and in semen, because irregularly shaped cells in the seminal duct could be either parasperm or normal spermatids. Although parasperm are identified by clear criteria in vertebrates, only in cottoid fishes to date, it is possible for parasperm to be discovered in other vertebrates. Recently, roles related to sperm competition have been reported in several species (e.g., the marine cottoid fish Hemilepidotus gilberti), and, with some of them, parasperm production is influenced by an intraspecific factor such as a sex ratio or the density of a population. Sperm competition is one of the important candidates for influencing evolution of parasperm functions, but not all parasperm seem to have a relation to sperm competition. Parasperm function may relate to the ecological conditions of each species that produces parasperm. Studies on parasperm function will be advanced by an ecological approach concerning male fertilization success as well as cytological investigation for parasperm. An erratum to this article is available at .     

Ectaquasperm-like parasperm in an internally fertilizing gastropod

Abstract. Pomacea canaliculata is an internally fertilizing gastropod that produces, besides fertilizing sperm (=eusperm), a large number of unfertile sperm (=parasperm) that have no chromatin, are fusiform, and have three to five flagella. Here, we report that this snail also produces another type of parasperm, which results from a peculiar spermiogenesis including an anterior cytoplasmic migration. The mature oligopyrene parasperm has: (1) a rounded head including a partly lysed nucleus, (2) a conical mid-piece with eight large mitochondrial structures, and (3) a single flagellum (∼20 μm). These characteristics, although not found in any other gastropod parasperm, are shared with the externally fertilizing "ectaquasperm" and with the early spermiogenic stages of internally fertilizing "introsperm" found among the Annelida and basal Mollusca. There are indications that this sperm type may be produced by a truncation of euspermiogenesis, as proposed by Buckland-Nicks & Scheltema for the expression of ectaquasperm in bilaterian evolution.     

Characterization of an alternative chromatin remodeling to parasperm in a cottid fish, Hemilepidotus gilberti

The dimorphic sperm of Hemilepidotus gilberti, i.e., haploid eusperm and diploid parasperm, have different morphologies corresponding to their own roles in fertilization. To estimate how these specific sperm morphologies were established, we focused on the nuclear morphologies and examined their changing processes in dimorphic spermiogenesis. Electron microscopic observation revealed that, in euspermatids, chromatin condensation first appeared as a mosaic pattern of moderate electrodense material in the peripheral region of the round nucleus. Those materials spread across the whole area to form a uniformly condensed nucleus. Chromatin condensation began similarly in paraspermatids to that in euspermatids. These became localized to one side of a nucleus and further condensed to form strong electrodense chromatin clusters, which are a specific feature of parasperm. From the remodeled nuclei of eusperm and parasperm, we found five and three kinds of sperm-specific basic proteins (SBPs), respectively, substituted to histones. The N-terminus amino acid sequences of the SBPs suggest that, in parasperm, one major SBP and two minor ones were distinct from each other. In eusperm nuclei, two kinds of specific SBPs were detected in addition to the homologs of parasperm SBPs. The specific SBPs had homologous amino acid sequences with huge arginine clusters, and one of them was most dominant among the five kinds of SBPs. The different combinations of SBPs in the eusperm and parasperm may cause a specific pattern of chromatin condensation in the dimorphic sperm nuclei of H. gilberti.     

Nuclear fragmentation characterises paraspermiogenesis in Tubifex tubifex (Annelida, Oligochaeta)

It is known that tubificine oligochaetes produce two types of spermatozoa: eusperm, fertilizing sperm with regular haploid DNA content; and parasperm, with a much lower DNA content, protecting and carrying the eusperm. Whereas mature spermatozoa and spermatids of the two lines are easily recognized by their morphology and DNA content, little is known about the first steps of differentiation of the two lines. This subject is addressed here in two ways: we have measured DNA content by a new method based on confocal laser microscopy and found that the total DNA content of parasperm cysts is extremely variable and equal or lower than total DNA content of eusperm cysts. Then we focused on the spermatocytes, and we found that the cells which will form paraspermatids undergo a peculiar kind of nuclear fragmentation which differ greatly from a regular cell division. During fragmentation DNA is distributed unevenly among the spermatids and this gives rise to a great and variable number of parasperm with variable DNA content. Immunocytochemical assays revealed that a proper meiotic spindle is never formed during fragmentation and that actin may play an important role in the chromatin division. Electron micrographs showed that the centrioles undergo a phenomenon of mass reproduction similar to that found in ciliated epithelia which supplies each of the numerous paraspermatids of its basal body. Mol. Reprod. Dev. 59: 442-450, 2001.     

Obstructive Role of the Dimorphic Sperm in a Non-copulatory Marine Sculpin, Hemilepidotus gilberti, to Prevent Other Males' Eusperm from Fertilization

In many species of animals, males normally produce parasperm (dimorphic sperm) along with eusperm (normal sperm) during spermatogenesis. In the present study, to clarify the role of parasperm of the non-copulatory sculpin Hemilepidotus gilberti, whose reproduction is characterized by polyandrous oviposition involving sneaking by neighboring territorial males, we observed the movements of parasperm. Parasperm could not move by themselves, but they were transported in solutions by passive movement due to collisions with actively swimming eusperm. In the viscous ovarian fluid (OF), which isolates eggs from seawater by covering them during spawning, parasperm did not exhibit any movement. However, they could be transported by eusperm movement in solutions with dissolved OF, partly because the viscosity of the fluid become lower. And then, in some solutions parasperm formed lumps. Lump formation of parasperm was also observed at the boundary surface of an egg mass where OF contacted seawater. Eusperm added experimentally to a solution in which parasperm were forming lumps were engulfed in the lumps and never escaped. Thus, lump formation of parasperm would be obstacles for the later arriving eusperm. Although lumps formed against both kin and non-kin eusperm, parasperm are thought to be available to overcome sperm competition which would occur during spawning that involves sneaking being almost concurrent with lump formation. The territorial male eusperm reach the eggs while his parasperm hinder other males' eusperm from reaching the eggs. Thus, we concluded that parasperm of H. gilberti play a role on protection of paternity by blocking rival eusperm physically from approaching eggs.     

Pseudopollen: Its Structure and Development in Maxillaria(Orchidaceae)

Histochemical analyses of the pseudopollen of ten species ofMaxillaria sectionGrandiflorae revealed that the main storageproduct is protein, although starch is usually also present.Lipids are rare in pseudopollen and thus do not seem to playan important role in attracting insects. In Maxillaria sanderiana,pseudopollen is formed by the fragmentation of multicellular,uniseriate trichomes, derived by the repeated division of asingle, papilla-like, basal secretory cell that contains well-developeddictyosomes, endoplasmic reticulum and mitochondria. At first,there is continuity of cytoplasm between adjacent componentcells of a trichome via plasmodesmata. During maturation, thecytoplasm retracts as the cell volume increases and the plasmodesmatabecome less obvious. Each component cell of the trichome eventuallycomprises a large protein body and a small amount of peripheralcytoplasm containing amyloplasts, a few small lipid bodies,mitochondria and a nucleus with nucleolus. Finally, the trichomeundergoes fragmentation, forming individual cells or chainsof cells of varying lengths. Light microscopy observations indicatea similar sequence in the other species examined. The occurrenceof pseudopollen in section Grandiflorae and alliance Splendensmay indicate that this character has evolved at least twicein Maxillaria. Copyright 2000 Annals of Botany Company Bees, farina, histochemistry, labellum, low-vacuum scanning electron microscopy, Maxillaria, Orchidaceae, pseudopollen, transmission electron microscopy, trichomes     

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)     

Ultrastructural Aspects of the Glandular Cells from the Secretory Trichomes and from the Cell Suspension Cultures of Achillea millefolium L. ssp. millefolium

The ultrastructure of the glandular cells of the floret secretorytrichomes from Achillea millefolium L. ssp. millefolium (yarrow)was examined before and after anthesis and compared with theultrastructure of the cells from the cell suspension culturesobtained from the same plant. The profuse tubular structuresobserved in the plastids of the glandular cells of the trichomesduring the pre-secretory stage were much reduced in the secretorystage and showed an osmiophilic content. Some endoplasmic reticulumprofiles could be seen adjacent to the plastids. Later in thesecretory stage, the secretion appeared in the periplasmic spacebetween the cells of the upper tiers and in the sub-cuticularspace. Finally the secretion was released by rupture of thecuticle. At the lag phase, the cells from the cell suspensioncultures of yarrow were characterized by the presence or plastidswith tubular structures, similar to those observed in the plastidsof the trichomes in the pre-secretory stage. By the end of thelag phase accumulations of starch were observed inside the plastids.At the beginning of the exponential phase, the tubular structuresof the plastids started to show an osmiophilic content and theaccumulations of starch were still present. At the end of thisphase starch disappeared from the plastids and only osmiophilictubular structures were observed. Rough endoplasmic reticulumas well as smooth endoplasmic reticulum profiles were frequentlyin close association with plastids and mitochondria. At thestationary phase a very large vacuole filled the cells, andin the remaining cytoplasm some endoplasmic reticulum profilesand osmiophilic droplets were observed.Copyright 1994, 1999Academic Press Achillea millefolium L. spp. millefolium, yarrow, ultrastructure, trichomes, glandular cells, plant cell suspension cultures     

HISTOCHEMISTRY AND ULTRASTRUCTURE OF THE CRYPT CELLS IN THE DIGESTIVE GLAND OF APLYSIA PUNCTATA (CUVIER, 1803)

The crypt cells lining the Aplysia punctata digestive tubulescomprise of three types of cell; calcium, excretory, and thincells. The calcium cells play a role in osmoregulation, mineral storage,exocrine secretion, iron detoxification, and excretion processes.They possess well- developed microvilli and a basal labyrinth,suggesting a role in absorption. The Golgi apparatus is involvedin the production of two main components of calcium spherules;the fibrillar material and mineralized granules. Golgi complex,rough endoplasmic reticulum (RER), ribosomes, and altered mitochondriaare involved in the formation of calcium spherules. Secretoryactivity is indicated by the formation of dense granules containingiron and calcium salts. Lipofuscin pigment has been found inlarge concretions which may arise from cytoplasmic areas surrounded byendoplasmic reticulum, RER and Golgi tubules. There are threestages of excretory cells, called early, mature, and post-excretorycells. This study traces the development of granulofibrillarvacuoles up to the formation of the lipofuscin concretions andshows that excretory cells are in fact degenerating calciumcells. The fine structure of thin cells suggests that they areyoung calcium cells. (Received 29 December 1997; accepted 15 November 1998)     

Chromatin subunit structure in different tissues of higher plants

The basic chromatin structure of higher plants (Vicia faba andTrillium kamtschaticum) was examined biochemically. After digestionwith micrococcal nuclease, the chromatins of these species yieldedDNA-protein components which sedimented as discrete peaks at11S, 15S, 19S, and so on in a sucrose gradient. The buoyantdensity of Vicia chromatin subunits was about 1.44 g?cm^–3in CsCl. Polyacrylamide gel electrophoresis of histone fromthese subunits of Vicia and Trillium chromatins indicated thatthe 11S monomer contained very little histone H1 but a fullcomplement of all other histones, whereas the oligomers containedH1 as in the case of undigested chromatin. Therefore, the modeof organization of basic chromatin structure in higher plantsis identical with that reported with various other eukasyotes,although two plant histone components are different from thecorresponding mammalian histones, H2A and H2B, in molecularweight and amino acid composition. The results indicated alsothat chromosomes prepared from Trillium meiotic cells do notdiffer from chromatins of Trillium or Vicia somatic cells inthe sensitivity to nuclease digestion or in the size of theirsubunits. (Received May 19, 1978; )     

Ultrastructure of Nectaries of Vinca rosea L., Vinca major L. and Citrus sinensis Osbeck cv. Valencia and its Relation to the Mechanism of Nectar Secretion

The ultrastructural changes in nectar-secreting cells of Vincarosea, Vinca major, and Citrus sinensis during their ontogeneticdevelopment are described. The most pronounced changes occurin the amount and morphology of the endoplasmic reticulum (ER).The amount of ER elements increases gradually and reaches amaximum at the stage of secretion. At this stage the ER is thedominant element in the cytoplasm. A process of swelling ofa lamellar ER, followed by formation of vesicles, was notedin the secretory cells during the stage of secretion. Many vesicularelements appeared to be in a close association with the plasmalemma.It is suggested that sugar is secreted as a solution by meansof vesicles derived from the ER.     

THE PALLIAL EYES OF CTENOIDES FLORIDANUS (BIVALVIA: LIMOIDEA)

The structure of the pallial eye in the Limidae has neverbeen elucidated properly, largely because they are difficultto see among the mass of surrounding mantle tentacles and becausethey are few, small, and lose their pigmentation when preserved.Possibly two eye types are present, simple cup-shaped receptorsin species of Lima, like those seen in the Arcoida, and morecomplex invaginated ones in Ctenoides. The pallial eyes (;18on both lobes) of Ctenoides floridanus are formed by invaginationof the middle mantle fold at the periostracal groove, so thatall its contained structures are derived from the outer andlight is perceived through the inner epithelia of this fold.The eye comprises a simple multicellular lens and a photoreceptiveepithelium beneath it of lightly pigmented cells and alternatingvacuolated, support cells. In some species of the Arcoidea, Limopsoidea and Pterioidea, pallialeyes occur on the outer mantle fold and thus beneath the periostracum(and shell). The pallial eyes of Ctenoides floridanus and otherpterioideans, e.g. species of the Pectinidae, occur on the middlefold and may thus have improved vision. In the Cardiodea, Tridacniidaeand Laternulidae (Anomalodesmata) pallial eyes occur on theinner folds. There is thus a loose phylogenetic trend, in which Ctenoidesis a critical link, of increasing eye sophistication correlatedwith the historical age of the clades possessing them. (Received 16 November 1999; accepted 20 January 2000)     

Studies on the Sporogenous Lineage in the Moss Timmiella barbuloides IX. Development of the Tapetum

The ultrastructure of tapetal cells in Timmiela barbuloideswas investigated in relation to events of sporogenesis. Aftertheir establishment both internally and externally to the sporogonialinitials, tapetal cells enlarge and assume a permanently polarizedorganization after completion of meiosis. A large vacuole isformed in the cell region distal to the spore sac, the nucleusbecomes centrally located, and amyloplasts lie in the cytoplasmadjacent to the spore sac. An extensive endomembrane systemdevelops in tapetal cells during the stage of exine depositionin spore tetrads. Sheets of rough endoplasmic reticulum developfirst around the nucleus then also in close proximity to theplasma membrane abutting the spore sac. Concomitantly, interveningdictyosomes produce a variety of vesicles. Unusual structureswith vesicle-like profiles also occur in the inner tapetum cellwalls abutting the spore sac. At the same time most of the starchis lost from the plastids in which grana-fretwork systems develop.A massive secretion of extremely electron-opaque material isassociated with perine deposition onto the free spore surfaces.Degeneration of the tapetal cells during the terminal stagesof spore maturation is marked by distortion of the organelles,increase in vacuolation and the appearance of electron-opaquematerial between the sheets of endoplasmic reticulum.Copyright1994, 1999 Academic Press Bryophytes, endomembrane dynamics, Timmiella, ultrastructure, development, tapetum     

稻虱跗煽的个体发生研究：卵的形成和胚胎发育

捻翅目昆虫是胎生的,胚胎发育和孵化均在母体血腔内进行。稻虱跗蝇Elenchinus japonicus属捻翅目,跗煽科,寄生于白背稻虱、褐稻虱和灰稻虱。本文报道稻虱跗煽卵的形成各阶段：1)雌虫体内无典型的卵巢,所有卵在母体体腔内同步发育和成熟。最早发现的原卵是包囊干细胞,在雌幼虫血腔内。2)每个包囊细胞内含256个姐妹细胞,其中有一个细胞分化成卵母细胞,其余的成为营养细胞。3)成熟卵为椭圆形,大小为95-100X40-50μm。其胚胎发育过程按顺序包括：卵裂、胎盘形成、胚带分节、附肢形成和胚胎背合等阶段。稻虱跗蛔行单胚生殖。     

苦皮藤素V对东方粘虫中肠细胞及其消化酶活性的影响

苦皮藤素V是从杀虫植物苦皮藤Celastrus angulatus Max.根皮中分离的一种对昆虫具有毒杀活性的新化合物。该文通过电镜观察和生化分析研究了其对东方粘虫Mythimnaseparata(walker)幼虫中肠组织及中肠主要消化酶活性的影响。电镜观察发现,中毒试虫的中肠细胞及其细胞器发生明显病变：柱状细胞顶膜微绒毛零乱、减少;线粒体肿胀,出现空白亮区,双层膜不完整;细胞质密度降低,细胞器排列紊乱;内质网池扩张,囊泡化,粗面内质网减少;杯状细胞杯腔变大,微绒毛减少。消化酶活性测定结果表明,中毒试虫中肠的蛋白酶、淀粉酶及脂肪酶的活性和正常虫相比,无显著变化。因此认为,苦皮藤素V主要作用于中肠细胞的质膜及其内膜系统。     

Dimorphic Sperm Influence Semen Distribution in a Non-copulatory Sculpin Hemilepidotus Gilberti

We used an artificial semen emission test to examine the semen transporting role of parasperm (unflagellated sperm), which are produced along with eusperm (normal sperm) by an incomplete meiosis in the marine sculpin Hemilepidotus gilberti. After separation of contents of semen (eusperm, parasperm, and seminal plasma) by centrifugation, the distance traveled by semen discharged vertically from a syringe into seawater was compared among various test semen; eusperm semen (eusperm re-mixed with seminal plasma), parasperm semen (parasperm re-mixed with seminal plasma), normal semen (eusperm and parasperm re-mixed with seminal plasma), and natural semen (unmodified semen). Parasperm semen traveled more than 1.5 times as far as the eusperm semen. The lateral dispersion width of normal semen after emission was significantly narrower than that of eusperm semen. These findings indicate that parasperm can reduce the lateral dispersion and prolong the distance semen travels. The eusperm ratio in the lower portion of the emitted semen did not differ from that in the upper portion, indicating that eusperm evenly distribute within the ejaculate and reach the lower portion of semen. Since males cannot closely approach eggs that are deposited in the narrow space between the female's belly and the spawning substrate, restraint of lateral dispersion and prolongation of the distance semen traveled would increase the number of eusperm arriving at eggs, despite reduction of eusperm in the ejaculate.     

The Digestive Glands of Pinguicula: Structure and Cytochemistry

The digestive glands of the carnivorous genus Pinguicula havethree functional compartments, (a) a basal reservoir cell, (b)an intervening cell of endodermal character and (c) a groupof secretory head cells. The gland complex is derived from asingle epidermal initial. The reservoir cell, which is richin Cl^– ions, is highly turgid before discharge; it islinked by plasmodesmata to the surrounding epidermal cells,and is ensheathed by a pectin-rich inner wall layer. The endodermalcell is bounded by a Casparian strip to which the plasmalemmais tightly attached; it contains abundant storage lipid andnumerous mitochondria. The head cells of the developing glandhave labyrinthine radial walls of the transfer-cell type, theingrowths being composed of pectic polysaccharides. The boundingcuticle is discontinuous, although lacking well-formed pores.Mitochondria are numerous, with well-developed cristae; theplastids are large and ramifying, and invested by ribosomalendoplasmic reticulum. Dictyosomes are sparse, and where theyoccur, are associated with coated vesicles. Ribosomal endoplasmicreticulum is moderately abundant in the head cells, and so alsoare free ribosomes. Optical and electron microscopic localizationmethods indicate that the digestive enzymes are synthesizedin the head cells and transferred both into the vacuoles andinto the walls. There is no evidence of a granulocrine modeof secretion, and the transfer seems to be initially by directperfusion through the plasmalemma. During the final phase ofmaturation of the head cells they suffer a form of autolysis,vacuoles, cytoplasm and wall becoming confluent as all of themembranes of the cell undergo dissolution. The gland head isthus, in effect, simply a sac of enzymes at the time of theultimate discharge. Pinguicula, carnivorous plant, insectivorous plant, enzyme secretion, digestive gland