Chiton integument: Ultrastructure of the sensory hairs of Mopalia muscosa (Mollusca: Polyplacophora)

Summary The dorsal integument of the girdle of the chiton Mopalia muscosa is covered by a chitinous cuticle about 0.1 mm in thickness. Within the cuticle are fusiform spicules composed of a central mass of pigment granules surrounded by a layer of calcium carbonate crystals. Tapered, curved chitinous hairs with a groove on the mesial surface pass through the cuticle and protrude above the surface. The spicules are produced by specialized groups of epidermal cells called spiniferous papillae and the hairs are produced by trichogenous papillae. Processes of pigment cells containing green granules are scattered among the cells of each type of papilla and among the common epidermal cells.The wall or cortex of each hair is composed of two layers. The cortex surrounds a central medulla that contains matrix material of low density and from 1 to 20 axial bundles of dendrites. The number of bundles within the medulla varies with the size of the hair. Each bundle contains from 1 to 25 dendrites ensheathed by processes of supporting cells. The dendrites and supporting sheath arise from epidermal cells of the central part of the papilla. At the base of each trichogenous papilla are several nerves that pass into the dermis. Two questions remain unresolved. The function of the hairs is unknown, and we have not determined whether the sensory cells are primary sensory neurons or secondary sensory cells.     

Chiton integument: development of sensory organs in juvenile Mopalia muscosa

The girdle epidermis of adult Mopalia muscosa secretes several types of structures, including calcareous spicules and innervated hairs. Newly metamorphosed chitons superficially resemble adult animals, but they lack the adult girdle ornaments, shell sculpture, and coloration. The morphogenesis of the adult girdle structures has not been described previously for any species. Juvenile Mopalia muscosa secrete hairs at metamorphosis, but it was not known if these hairs were sensory or if they were retained as the animals grew. I discovered that the hairs of juveniles become the tips of adult hairs. When juvenile hairs are detectable by light microscopy the sensory components already exist, suggesting that they are functional receptor organs. The other girdle ornaments of young juveniles, the primary calcareous spicules, are lost as the animal grows. I also demonstrated that the hairs are not uniquely innervated; the same sensory structures are produced in conjunction with other girdle ornaments on the marginal and ventral faces.     

The chiton gill: Ultrastructure in Chiton olivaceus (Mollusca,Polyplacophora)

Gills of Chiton olivaceus, a primitive mollusc, are relatively simple in their structure and ultrastructure but are well adapted to a life in the intertidal zone. In contrast to some other molluscs, there is no differentiation of the gill epithelium into functional regions other than respiratory ones. Ciliation of the epithelium in certain areas may optimize water flow from the outer to the inner part of the mantle cavity. The hemolymph sinuses are oriented so that hemolymph flows in the opposite direction. Interstitial cells link epithelial cells with nerve endings. Muscle cells as well as the collagenous matrix in the connective tissue differ within the main gill axis and the lateral lamellae. The life cycle of immunoactive cells within the connective tissue and the hemolymph is described.     

Neurogenesis in the mossy chiton,Mopalia muscosa (Gould) (Polyplacophora): evidence against molluscan metamerism

Neurogenesis in the chiton Mopalia muscosa (Gould, 1846) was investigated by applying differential interference contrast microscopy, semithin serial sectioning combined with reconstruction techniques, as well as confocal laser scanning microscopy for the detection of fluorescence-conjugated antibodies against serotonin and FMRFamide. The ontogeny of serotonergic nervous structures starts with cells of the apical organ followed by those of the cerebral commissure, whereas the serotonergic prototroch innervation, pedal system, and the lateral cords develop later. In addition, there are eight symmetrically arranged serotonergic sensory cells in the dorsal pretrochal area of the larva. FMRFamide-positive neural elements include the cerebral commissure, specific "ampullary" sensory cells in the pretrochal region, as well as the larval lateral and pedal system. In the early juvenile the cerebral system no longer stains with either of the two antibodies and the pedal system lacks anti-FMRFamide immunoreactivity. Outgroup comparison with all other molluscan classes and related phyla suggests that the cord-like, nonganglionized cerebral system in the Polyplacophora is a reduced condition rather than a primitive molluscan condition. The immunosensitivity of the pedal commissures develops from posterior to anterior, suggesting independent serial repetition rather than annelid-like conditions and there is no trace of true segmentation during nervous system development. Polyplacophoran neurogenesis and all other available data on the subject contradict the idea of a segmented molluscan stem species.     

Preservation potential of Katharina tunicata and Mopalia muscosa (Mollusca, Polyplacophora) on two rocky shores of San Juan Island, Washington, USA

Taphonomic analyses of modern Katharina tunicata and Mopalia muscosa valves (Mollusca, Polyplacophora) collected from San Juan Island, Washington, USA, demonstrate that preservation potential of chitons varies by species and locality. Damage levels observed in the valves reflect differences in extrinsic environmental conditions, but intrinsic characters affect response of the skeletal material to local processes. Although both are more likely to become fossils at the lower energy site, K. tunicata valves in poor condition are more likely to occur than poor-condition M. muscosa valves at either site. Mopalia muscosa valves are least likely to be preserved at the higher energy site. Katharina tunicata valves appear more resistant to destruction than M. muscosa valves, suggesting that the former are more likely to become fossilized. The preservation potential of the three types of valves (normally one head, six medials, and one tail per individual) differs for both species. Medials are more likely to be preserved than either terminal valve of K. tunicata or M. muscosa . Head valves are least likely to be preserved, but K. tunicata heads are less likely to be preserved than M. muscosa heads. The biases are not due to greater frequency of medials, because valves do not occur in the 3:1 ratio seen in living animals. Divergence from expected ratios and variation in taphonomic condition for all three valve types in modern environments agrees with observations in the fossil record. The rarity of fossil chitons may be due to biases against preservation rather than absence in ancient environments.     

Fine structure and immunocytochemistry of a new chemosensory system in the Chiton larva (Mollusca: Polyplacophora).

Combined electron microscopy and immunocytochemistry of the larvae of several polyplacophoran species (Chiton olivaceus, Lepidochitona aff. corrugata, Mopalia muscosa) revealed a sensory system new to science, a so-called "ampullary system." The cells of the "ampullary system" are arranged in four symmetrically situated pairs lying dorsolaterally and ventrolaterally in the pretrochal part of the trochophore-like larva and they send axons into the cerebral commissure. They are lost at metamorphosis. The fine structure of these cells strongly resembles that of so-called "ampullary cells" known from various sensory organs of other molluscs, such as the apical complex of gastropod and bivalve larvae, osphradia of vetigastropods, and olfactory organs of cephalopods, and nuchal organs of certain polychaetes. The ampullary cells and their nerves are densely stained by anti-FMRF-amide fluorescence dyes, whereas antiserotonin staining is only weak. While cytological homology of the ampullary cells with those of other organs is probable, the ampullary system as a whole is regarded as a synapomorphy of the Polyplacophora or Chitonida.     

Reproductive traits and relative gonad expenditure of the sexes of the free spawning Chiton articulatus (Mollusca: Polyplacophora)

ABSTRACT

Reproductive studies of an intertidal free-spawning population of Chiton articulatus (Mollusca: Polyplacophora) from Puerto Angel, Oaxaca, Mexico were undertaken during 2011. We used gonad histology and gonadal indices to assess the relative gonad expenditure of the sexes (RGES) and other reproductive traits, accounting for individual and seasonal variation within this population. At this location, C. articulatus is gonochoric, without sexual dimorphism, except internally by gonad colour (testis is ‘salmon’ coloured and ovary olive green). Annual and monthly sex ratios (m/f) do not differ significantly from 1:1. Highest population-level gonadosomatic index (GSI) corresponded to maximum (peak) ripe stage (i.e. maximum gonad investment), with a first peak in May with a high value (8.4 ± 0.5) and a second peak during August-September with a lower value (4.7 ± 0.3). GSI fluctuated throughout year implying that gonad expenditure may be seasonally constrained, but with overall synchrony between sexes of ripe and spawning stages. July to December was the main reproductive season with some facultative spawning occurring off-season. Ripe and spawning RGES did not differ between sexes, suggesting that either sperm competition is intense and/or that sperm limitation is high. Early spawning individuals may quickly replenish their gametes for a second phase of gamete release later.

Abbreviations: RGES: relative gonad expenditure of the sexes; GSI: gonadosomatic index; GDS: gonad developmental stages; MiMI: microscopic maturity index; SST: sea-surface temperature.     

Hermaphroditism in two populations of Chiton articulatus (Mollusca: Polyplacophora) from the eastern tropical coast of México

Through histological analyses, this study reveals an unusually high incidence of hermaphroditism in Chiton articulatus. Specimens were sampled every 30?days between September 2010 and September 2011 at two locations (Las Brisas and Jaramillo beaches, Acapulco) on the tropical eastern Pacific coast of Mexico. At both sites, hermaphroditism was found throughout the year, although in varying proportions. Higher percentages of hermaphroditism were found during the pre-spawning summer months (Las Brisas Beach 63%, Jaramillo Beach 68%). Two different kinds of hermaphroditic gonads were found, showing a preponderance of either male or female tissues but, commonly, female tissues occupied the greatest part of the gonad cross sections. Similar to other species of polyplacophorans, there was a predominance of males, although M:F sex ratios ranged from 0.7 to 4.5: 1 at Las Brisas Beach and 0.3–8: 1 at Jaramillo Beach.     

Chitons (Mollusca: Polyplacophora) from El Salvador, Central America

Collections of 11 species of shallow water Polyplacophora from El Salvador were made in July 2002. Previously only five species had been documented in El Salvador: Chaetopleura lurida (Sowerby, 1832); Ischnochiton guatemalensis (Thiele, 1910); Ceratozona angusta (Thiele, 1909); Chiton stokesii (Broderip, 1832) and Acantochitona exquisita (Pilsbry, 1893). Of these, L. guatemalensis and A. exquisita were not collected in this census. Seven other species are reported here for El Salvador for the first time: Lepidochitona beanii (Carpenter, 1857); Ischnochiton dispar (Sowerby, 1832); Stenoplax limaciformis (Sowerby, 1832); Callistochiton expressus (Carpenter, 1865); Acanthochitona arragonites (Carpenter, 1867); A. ferreirai (Lyons, 1988) and A. hirudiniformis (Sowerby, 1832). The known geographic distribution of 1. dispar is extended to the north. An un-named species of Lepidochitona is briefly described.     

Observation and establishment of gonad development stages in polyplacophorans (Mollusca): Chiton (Chiton) articulatus a case study

Gonad development stages (GDS) and, subsequently, the reproductive cycle are described by performing histology of some gonad portions. In polyplacophorans, gametogenesis is not enough to define GDS; further anatomical gonad features are relevant. In most adult polyplacophorans, the gonad is a simple anatomical structure that resembles and operates as one single gonadal acinus without glandular structure. These features have gone unnoticed causing inaccurate GDS assignment and, consequently, imprecise reproductive season in polyplacophorans. Here, dissection protocols that allow extracting a compact gonad are established. Emphasizing the anatomical structure of the whole gonad and the displacement of gametes, five GDS were assigned to both sexes of Chiton (Chiton) articulatus: I‐goniogenesis, II‐development, III‐ripe, IV‐spawning, V‐rest. Tissue platelets contribute importantly to GDS assignment and even help distinguishing between males and females. Neither a randomly selected portion of gonad nor a longitudinal section are recommended because it leads to misinterpretation higher than 50% in determining GDS and besides ignores displacement of gametes. A panoramic sweep across a complete transverse‐section of each gonad was validated as the best option for establishing GDS. This new methodology was tested on several polyplacophorans species, and seems generally applicable for histological assessment of reproductive cycle and reproductive season in polyplacophorans.     

Spermiogenesis in Polyplacophora,with special reference to acrosome formation (Mollusca)

Summary The present study examines spermiogenesis, and in particular the formation of the acrosome, in ten species of chitons belonging to four families. This study emphasizes the formation of the acrosome but brings to light several other structures that have received little or no mention in previous studies. The process of spermiogenesis is essentially similar in each species, although Chaetopleura exhibits some significant differences. In early spermiogenesis the Golgi body secretes numerous small pro-acrosomal vesicles that gradually migrate into the apical cytoplasm. The chromatin condenses from granules into fibres which become twisted within the nucleus. A small bundle of chromatin fibres projects from the main nuclear mass into the anterior filament; this coincides with the appearance of a developing manchette of microtubules around the nucleus that originates from the two centrioles. Radiating from the distal centriole is the centriolar satellite complex, which is attached to the plasma membrane by the annulus. The distal centriole produces the flagellum posteriorly and it exits eccentrically through a ring of folded membrane that houses the annulus. Extending from the annulus on one side of the flagellum, in all but one species, is a dense fibrous body that has not been previously reported. The proximal centriole lies perpendicular to the end of the distal centriole and is attached to it by fibro-granular material. Pro-acrosomal vesicles migrate anteriorly through the cytoplasm and move into the anterior filament to one side of the expanding nucleus. Eventually these vesicles migrate all the way to the tip of the sperm, where they fuse to form one of two granules in the acrosome. In mature sperm the nucleus is bullet-shaped with a long anterior filament and contains dense chromatin with occasional lacunae. The mitochondria vary in both number and position in the mature sperm of different species. Both centrioles are housed eccentrically in a posterior indentation of the nucleus, where the membranes are modified. The elongate flagellum tapers to a long filamentous end-piece that roughly corresponds to the anterior filament and may be important in sperm locomotion for hydrodynamic reasons. An acrosome is present in all ten species and stained positively for acid phosphatase in three species that were tested.     

Die Feinstruktur der Ästheten vonChiton olivaceus (Mollusca,Polyplacophora)

The aesthete organs in the shell of the polyplacophoranChiton olivaceus (Spengler) were studied by scanning and transmission electron microscopy. Compared to previously described species they reveal marked differences. In the upper third of the aesthete, photoreceptor cells have been found. The granula of the club-shaped cells, which fill most of the aesthete, are formed in the proximal part of young aesthetes. The secretory cells located in this part increase in size and become gradually club-shaped. The central cells, hitherto the only known sensory cells in this organ, are very variable in their appearence. The apical as well as the subsidiary caps have pores which penetrate the whole cap in the former, whereas the subsidiary caps are proximally and distally covered by a continous envelope. In contrast, an exchange of substances with the surrounding environment cannot be excluded in the apical cap: there are some indications of secretory processes occurring through the shell surface.     

The valve morphology of Cullochhiton uchutinus (Mollusca: Polyplacophora: Ischnochitonidae)

The valves of Callochiton achatinus comprise four layers which are penetrated by channels carrying extensions of the mantle epithelium known as aesthetes. Three patterns of dorsal sculpturing within defined areas of the various valves are described and these are related to the different types of aesthete channel which perforate the respective areas. The processes involved in valve growth are described and related to the arrangement of the many holes penetrating the eave tissue.     

The Larval Development of the Chiton <Emphasis Type="Italic">Deshayesiella curvata</Emphasis> (Carpenter in Dall, 1879) (Mollusca: Polyplacophora)

Chronological study of the larval development of the chiton Deshayesiella curvata with sketching of the first stages of egg division and embryos up to the seven-valved shell stage is presented for the first time. D. curvata exhibits pelagic development with lecithotrophic larvae. Larval eyes form at the age of 3 days; at the age of 6 days, a Schwabe organ appears near the eyes in the form of two pigmented spots. Chitons at the stage of seven-valved shell have larval eyes, as well as the Schwabe organ.     

Die Mantelpapillen und Stacheln vonAcanthochiton fascicularis L. (Mollusca,Polyplacophora)

Zusammenfassung Struktur und Ultrastruktur der Papillen im Mantelepithel der PolyplacophoreAcanthochiton fascicularis werden beschrieben. Sie bestehen zu einem gro\en Teil aus Sekretzellen. In vielen Papillen sind Sehzellen vorhanden. Die Papillen bilden einen bis mehrere Stacheln aus. Ein organischer Becher verbindet den Kalkstachel mit einer Stachelzelle, hÄufig noch mit einer zweiten Zelle, die distal eine Cilie trÄgt. Sie sind Tastrezeptoren. Andere Stacheln dienen lediglich der Abwehr; sie können durch Muskeln bewegt werden. Diese Stacheln wachsen stÄndig basal nach, wÄhrend die Taststacheln nach einiger Zeit abgesto\en und durch neu von der Papille gebildete ersetzt werden. Ästheten und Mantelpapillen sind homologe Orgame.

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The mantle papillae and the spines inAcanthochiton fascicularis L. (Mollusca, Polyplacophora)

Summary Structure and ultrastructure of the papillae in the mantle epithelium of the polyplacophoranAcanthochiton fascicularis are described. They consist to a major part of various secretory cells. Visual cells occur frequently in the papillae. Each of these organs form up to a few spines which have basally a cup of organic material. This connects the calcareous spine with the spine cell and often with a second cell which has distally one cilium. They are tactile receptors. Other spines are only for the defense and can be moved by muscles. They continue to grow basally in contrast to the tactile spines which are pushed off after some time and replaced by new ones formed in the papillae. The aesthetes and the mantle papillae are homologous organs.

     

Serotonin-and FMRFamide-immunoreactive nerve elements in the chiton Lepidopleurus asellus (Mollusca,Polyplacophora)

The distribution of serotonin-like and FMRFamide-like immunoreactive (5HT-ir and FMRFa-ir, respectively) neurons in the nervous system of the chiton Lepidopleurus asellus (Mollusca, Polyplacophora) was studied using an immunocytochemical technique. The neurons were distributed in characteristic patterns in the central nervous system, the 5HT-ir neurons predominating in the ventral (pedal) cords and FMRFa-ir neurons in the lateral cords. In the body wall including the foot, a tight network of 5HT-ir and FMRFa-ir nerve fibers is found, the former being mostly attributed to the musculature whereas the latter seems to be associated with the blood sinuses. Intraepithelial neurons of both types are abundant in the fore-and hindgut. The presence and general distribution in the central and peripheral nervous system of the 5HT-ir and FMRFa-ir elements seems thus to be similar in simple and advanced molluscs. The relationship between these neurons and their targets in the body also appears to be well conserved in molluscs.     

Fine structure of the eyes of Onithochiton neglectus (Mollusca: Polyplacophora)

Summary Onithochiton neglectus a common littoral chiton possesses large numbers of small eyes embedded in the outer layer of the shell, the tegmentum. These are arranged in a definite pattern on each shell valve. Each eye lies in a pocket, and is surrounded by pigment laid down in the shell. There is a lens, cup of retina cells and an optic nerve running in an optic canal through the shell. Glial elements are present. The retina cells give rise centrally to a packed array of microvilli, a rhabdom. Cilia are present at the edge of the rhabdom; they have a 9 + 2 arrangement of ciliary filaments and do not appear to be involved in the formation of microvilli. Cells at the periphery of the eye cup give rise to large whorls of membranes, lamellate bodies. These bodies are derived from the membranes of cilia having a 9 + 2 pattern, and form into an extra-cellular space. Nerve processes from the retina cells pass into the optic canal. On the basis of previous work it is thought that the lamellate bodies are also sensory. These structures are discussed in relation to other microvillar and lamellate structures described from photoreceptors.I thank Professor J. E. Morton for his advice in the early stages of this work, and Dr. S. J. Bullivant for the fixation and embedding of material for electron microscopy. To Professor G. A. Horridge I am grateful for advice and the facilities of his laboratory, and to Professor M. S. Laverack, Patricia Holborow and Charles Coleman for much help and encouragement. I am supported by the Science Research Council, and in New Zealand held a Commonwealth Scholarship.     

Allometric and morphological characteristics of Tonicella marmorea (Fabricius, 1780) populations (Mollusca: Polyplacophora: Ischnochitonidae)

Allometric and morphological characteristics of population samples of Tonicella marmorea from three sea lochs on the west coast of Scotland are described. The suitability of allometric relationships as taxonomic criteria are investigated by statistical comparisons of various regression constants, scatter diagrams and comparisons of curvature indices. Morphological characteristics such as ctenidia number and numbers of notches in the head and tail valves proved too variable to be regarded as diagnostic but they are useful taxonomic indicators. The need for all taxonomic characteristics to be derived from population samples is stressed and the use of valve and girdle sculpturing together with radula structure is emphasized.     

Egg hull formation in Callochiton dentatus (Mollusca,Polyplacophora): the contribution of microapocrine secretion

Abstract. Egg hull formation during oogenesis in the chiton Callochiton dentatus does not follow the typical model of merocrine secretion involving Golgi vesicle exocytosis. Instead, microapocrine secretions are primarily responsible for egg hull formation, although merocrine secretions contribute “areolae” and the vitelline layer. Microapocrine secretion mechanisms are poorly understood, involving a different cellular pathway than is typical. Egg hull formation in C. dentatus involves two types of microapocrine secretions released by the oocyte, one of which is described here for the first time. The plesiomorphic jelly-like egg hull of chitons, as exemplified by the eggs of members of the basal order Lepidopleurida and present also in eggs of C. dentatus (Chitonida: Callichitonidae), may have evolved solely as an oocyte secretion, whereas members of some other families in the order Chitonida form their egg hulls with considerable secretory input from the follicle cells as well.     

Feinstrukturelle Untersuchungen zur Oogenese der KäferschneckeLepidochitona cinereus (Mollusca,Polyplacophora)

Zusammenfassung 1. Die Wachstumsphase der Oocyten vonLepidochitona cinereus L. läßt sich nach cytomorphologischen und histologischen Gesichtspunkten in fünf Entwicklungsstadien einteilen.2. Typische Oogonien in den Ovarien adulter Weibchen sind nicht eindeutig nachzuweisen. Es ist noch kein kugeliger Nucleolus ausgebildet; der Kern ist teilweise dicht mit Chromatinkörpern angefüllt. Diese Eizellen sind noch nicht vollständig durch Follikelzellen getrennt.3. Die Oocyten des Stadiums I sind stets von Follikelzellen umgeben. Ribosomenähnliche Grana akkumulieren sich zum kugeligen Nucleolus oder sind auf dem Wege zur Emission durch die Kernhülle. Sie bilden einen osmiophilen juxtanucleären Saum. Das Oocytoplasma enthält wenige kleine Mitochondrien, einen kleinen Golgi-Apparat, einige Lipidschollen und einzelne Stränge des Endoplasmatischen Retikulums.4. Charakteristisch für das Stadium II sind 1 bis 2 Dotterkerne (Balbiani-Körper), die sich aus einer dichten Wolke von Ribosomen und einer Schale aus Mitochondrien zusammensetzen. Der Nucleolus tritt in eine aktive Phase, zoniert sich in ein vakuoläres Zentrum und einen homogenen Cortex und schnürt von nun an Nucleolargrana (Paranucleoli) ab.5. Die vitellogenetische Phase wird im Entwicklungsstadium III durch die Vakuolisierung des Cytoplasma eingeleitet. Es können drei Vakuolentypen unterschieden werden, die ihren Ursprung entweder im Grundcytoplasma selbst, in Vesikeln des ER oder in benachbarten Golgi-Feldern haben können. Das ER entwickelt sich zu konzentrischen Membranstapeln.6. Das Stadium IV ist gekennzeichnet durch cytoplasmatische Aufwölbungen mit gleichzeitigen Einsenkungen der Oocytenmembran, die auf die Tätigkeit des oocytären Follikelepithels zurückzuführen sind. Es werden möglicherweise drei Wege der Dottersynthese beschritten: (a) In der inneren Mitochondrienmatrix akkumulieren sich Substanzen, die zu Protein-Primordialdotter mit Myelinlamellen führen. (b) Pinocytotische Vesikeln beteiligen sich an der Genese von homogen strukturierten Eiweiß-Dotterkugeln. (c) Lipidschollen gehen aus cytoplasmatischen Vakuolen hervor, die mit den umfangreichen Stapeln der annulate lamellae in Verbindung gestanden haben. Beim Übergang zum Stadium IV scheidet hauptsächlich die Oocyte über Mikrovilli eine aus Mucopolysacchariden bestehende dreischichtige, kompliziert gebaute Oocytenhülle ab. Im Cytoplasma liegen rootlets, die im Aufbau Cilienwurzeln gleichen.7. Im Stadium V erscheinen im cortikalen Cytoplasma Rindenvakuolen. Die Kernhülle faltet sich; die Nucleoli zerfallen. Die ausgereiften Protein-Dotterpartikeln enthalten ein homogenes Internum mit aufgelagertem parakristallin strukturiertem Cortex. Durch Anlagerung von Vesikeln und Lipidkörpern entsteht schließlich typischer Lipoproteindotter. Die Eizelle gibt eine weitere primäre Oocytenhülle, die Dottermembran ab.

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Ultrastructural investigations of oogenesis in the chiton,Lepidochitona cinereus (Mollusca, Polyplacophora)

Oocyte development ofLepidochitona cinereus L. has been examined by electron microscope with special regard to ultrastructural changes during vitellogenesis. Oogenesis can be subdivided into five stages based on cytological and histochemical features. Typical oogonia have not been found; only early oocytes of the meiotic prophase with an incomplete nucleolus are situated on the basallamina of the gonadal wall. The single nucleolus is formed in stage I; osmiophilic nucleolar granules pass through the nuclear envelope. Stage II — oocytes are characterized by simple Balbiani-bodies or yolk nuclei, which consist of ribosomes and a hull of mitochondria. The formerly homogenous nucleolus disintegrates into caryoplasmic vacuoles and produces paranucleoli. In the previtellogenetic stage III the yolk nuclei are reduced and large systems of endoplasmic reticula — arranged concentrically or flattened — and voluminous cytoplasmic vacuoles appear. Three types of vacuolar complexes can be observed: Complete membrane bounded vacuoles with a filaceous content built up by Golgi dictyosomes, vacuoles with remains of membranes which seem to originate from endoplasmic cisternae and vacuolar spaces lying free in the cytoplasm. The vacuoles contain either acid mucopolysaccharides or acid lipids. The actual vitellogenesis starts in stage IV after depression of the oocyte membrane to ooplasmic bumbs by the perioocytal follicle epithelium. Extensive piles of annulate lamellae contact the cytoplasmic vacuolar bodies. Striated long rootlets branch off microtubules at their terminal end in the direction of the oocyte membrane below the oocyte hull. Microvilli secrete mucopolysaccharides into the intercellular space between oocyte membrane and inner follicle cell membrane. The resulting eight cup-like hull processes are composed of three layers. Possibly there are three ways of forming vitelline bodies: (a) transforming mitochondria and multivesicular bodies lead to protein yolk; (b) mitochondria connect with cytoplasmic vacuoles and probably participate in genesis of lipid yolk; (c) microvesicles become protein yolk precursors. The lipoprotein yolk spheres consist of a homogenous internum and a paracrystalline structured cortex. Lipid yolk accumulates at the periphery of the cytoplasmic vacuoles, which degenerate later. The mature oocyte secretes another primary oocyte envelope, the vitelline membrane, shortly before spawning. Cortical granules appear below the oocyte membrane.