A new model for periostracum and shell formation in Unionidae (Bivalvia, Mollusca)

The periostracum in Unionidae consists of two layers. The outer one is secreted within the periostracal groove, while the inner layer is secreted by the epithelium of the outer mantle fold. The periostracum reaches its maximum thickness at the shell edge, where it reflects onto the shell surface. Biomineralization begins within the inner periostracum as fibrous spheruliths, which grow towards the shell interior, coalesce and compete mutually, originating the aragonitic outer prismatic shell layer. Prisms are fibrous polycrystalline aggregates. Internal growth lines indicate that their growth front is limited by the mantle surface. Transition to nacre is gradual. The first nacreous tablets grow by epitaxy onto the distal ends of prism fibres. Later growth proceeds onto previously deposited tablets. Our model involves two alternative stages. During active shell secretion, the mantle edge extends to fill the extrapallial space and the periostracal conveyor belt switches on, with the consequential secretion of periostracum and shell. During periods of inactivity, only the outer periostracum is secreted; this forms folds at the exit of the periostracal groove, leaving high-rank growth lines. Layers of inner periostracum are added occasionally to the shell interior during prolonged periods of inactivity in which the mantle is retracted.     

Histological description of the ink gland of the tropical sea hare Aplysia dactylomela Rang, 1828

This work describes for the first time the ink gland of Aplysia dactylomela Rang, 1828 using light microscopy and histochemical tests. The results reveal that this organ is covered by a layer of simple epithelium and that there are some differences between the epithelium facing the mantle shelf and that facing the mantle cavity. The former consists of columnar cells, which may have a secretory function, whereas the latter is a cuboidal epithelium. Underneath the epithelium and along the whole gland there are fibres of smooth muscle and collagen, organized in groups of parallel bundles. There are vesicles of different diameters, apparently with similar morphology. Some are filled with ink, whereas others are either granular or clear. The ink is released through a duct formed by invaginations of the cuboidal epithelium. Tests with bromophenol blue and periodic acid Schiff indicated that the ink and granulated vesicles contain proteins and carbohydrates or maybe glycoproteins. Between the fibre bundles and the vesicles there are dispersed cells. A diagram is presented emphasizing the covering epithelium, the distribution of muscle and collagen fibres, the dispersed cells, vesicles and ducts. This organization is similar to that of the other Aplysia species studied to date, A. californica.     

A HISTORY OF SKELETAL SECRETION AMONG ARTICULATE BRACHIOPODS

Studies of the ultrastructure of the exoskeleton of Notosaria nigricans (Sowerby), which can be used as the standard succession for articulate brachiopods, show that shell secretion involves six distinct operations giving rise to the following layers: mucopolysaccharide, outer fibrillar triple-layered membrane, mucoprotein, inner fibrillar triple-layered membrane, calcareous primary layer and calcareous-organic secondary layer. Comparison with the secretory régimes of terebratulids like Waltonia inconspicua (Sowerby) suggests that only four of these operations are fundamental, those controlling secretion of mucopolysaccharide, outer fibrillar membrane and the primary and secondary shell. An endoskeletal secretory phase giving rise to spicules in the mantle and lophophore is also found in many terebratulids. In the geological record, the orthodox primary-secondary shell can be traced back to the billingsellaceans and is presumed to have been associated with the mucopolysaccharide layer and a fibrillar membrane which together might well have constituted the prototypic organic exoskeleton. Deviation from this pattern during brachiopod evolution was relatively minor and was, so far as is known, limited to the strophomenids, some spiriferids and thecideidines.     

Ultrastructure of the pericardium in chitons (Mollusca: Polyplacophora), in relation to filtration and contraction mechanisms

Summary The pericardium in Lepidopleurus asellus (Spengler), Tonicella marmorea (Fabricius), T. rubra L., Ischnochiton albus L., and Calleochiton laevis (Montagu), species taxonomically far apart, is described. It consists of a flat, simple epithelium facing the pericardial cavity, a basement membrane, a muscle layer with two types of muscle fibres, nerve processes, glio-interstitial cells, and fibrocytes, embedded in a loose collagen matrix. The epithelium in L. asellus and I. albus have convoluted lateral cell borders, and in L. asellus very long basal cell processes are seen. Type 1 muscle fibres resemble smooth molluscan muscle. Type 2 muscle fibres resemble cardiac muscle fibres in chitons. Nerve processes associated with glio-interstitial cells and cell processes, run free in the matrix. Synapses in type 1 fibres are covered with glio-interstitial cell processes, lacking in type 2 muscle fibres synapses.This work was supported by grants from the Norwegian Research Council for Science and the Humanities     

Ultrastructure of the secondary tympanic membrane in the human fetus

The normal secondary tympanic membrane in human fetuses was examined by transmission electron microscopy. The membranes in 5- to 9-month-old fetuses consist of the following three layers: (1) an outer squamous epithelial layer facing the middle ear, which is not formed until 4 months old; (2) a middle fibrous layer containing collagen, elastin, fibroblasts and fibrocytes, and (3) an inner layer of flat cells facing the scala tympani. Following the maturation of the fetus the epithelium is getting thinner and fibroblasts are reduced in number, but fibrocytes are increased and collagen and elastin grow gradually in density. The ultrastructure of the secondary tympanic membrane at 8 month is mature in type and shows the same characteristics as in the adult. This membrane has an important and complicated physiological function. The epithelium of the outer layer, with tight junctions and multiple desmosomes, provides a barrier to keep harmful substances out. The stability of the membrane provides protection against rupture, while the elasticity plays a role in the physiology of hearing as well.     

Electron microscopic studies of growth margins of articulate brachiopods

Summary The brachiopod shell is secreted by the mantle epithelium lining the internal surfaces of its two valves. Growth lines, seen on their external surfaces, have been interpreted in terms of mantle regression and transgression from the valve margins. This scanning electron microscope study of the shell microstructure in recent brachiopods confirms these views and shows the skeletal evidence upon which such interpretations can be made. Electron micrographs reveal that from a growth line a plane dips posteriorly into the shell substance along which normal skeletal secretion was interrupted. Commonly a mosaic of secondary fibres, similar to that seen on the inside surface of the valve, is preserved upon this regression plane, most of the inside surface of which is covered by primary shell, usually extending posteriorly well into the secondary shell layer. The regression plane marks the area from which the mantle withdrew and the area over which shell secretion was interrupted. During mantle transgression primary shell was deposited over much of this surface, before the redevelopment of secondary fibres, so that the old internal surface of the valve was preserved as a false mosaic within the shell. In this way it is possible to recognise the extent of mantle regression and to note the position of the primary — secondary shell secreting junction of the mantle at the time when shell secretion was resumed.     

Morphology and structural organization of Gene''s organ in Argas walkerae

Electron microscopy revealed that Gene's organ in females of Argas walkerae Kaiser & Hoogstraal (Ixodida: Argasidae) is formed as a double-sac structure consisting of an outer epithelial and an inner cuticular sac. The latter emerges through the camerostomal aperture to the exterior in ovipositing ticks. The epithelial sac forms the corpus and the two blind-ending horns, which pass into the epithelium of the excretory duct of a gland at each side of Gene's organ and envelop the cuticular sac. Both excretory ducts open into the lumen between the epithelial and the cuticular sac. The cuticular sac is folded and consists of a fibrous endocuticula outwards towards the lumen between the epithelial and the cuticular sac and of a smooth epicuticula inwards. Parallel running grooves occur over the lateral epicuticular surface turning medially into cobble-stone pavement-like rises. Tubuli pass through the cuticular sac ending in pores on the epicuticular surface and open into the lumen between the epithelial and the cuticular sac. Muscle fibres pass through the epithelial sac at the horn tips and are inserted to the cuticular sac. In ovipositing females, the glands are fully developed and the lumen between the epithelial and the cuticular sac is filled with an amorphous mass.     

黄胫小车蝗受精囊的亚显微结构

利用组织学方法,观察了黄胫小车蝗Oedaleus infernalis 受精囊的显微与亚显微结构。结果表明,黄胫小车蝗受精囊为单个,由高度卷曲的受精囊管和蚕豆状的端囊构成。受精囊壁主要由表皮层、上皮层、基膜和肌肉层构成;上皮层包含上皮细胞、导管细胞和腺细胞。上皮细胞在靠表皮层的边缘有大量的微绒毛,两相邻上皮细胞的细胞膜相互嵌入,并有细微的突起延伸在导管细胞及腺细胞之间,直到基膜,达基膜处的上皮细胞膜折叠,与腺细胞膜的折叠,一起形成迷宫样的指状突起,附着在基膜上。导管细胞有一个较大的核和分泌导管,连接于腺细胞的细胞腔和受精囊腔,将腺细胞中分泌物运输到受精囊腔中。腺细胞具有典型的分泌细胞特征： 含发达内质网、高尔基复合体及不同大小的囊泡。肌肉层位于受精囊最外层,附在基膜上。在受精囊不同部位的结构有差异。在交配前和交配后,受精囊腺细胞的亚显微结构也有差异。     

The ultrastructure of the midgut and the formation of peritrophic membranes in a harvestman,Phalangium opilio (Chelicerata Phalangida)

Summary The ultrastructure of the midgut epithelium of Phalangium opilio was examined. In the anterior part of the midgut the epithelium consists of three different types of cells, called resorption, digestion, and excretion cells according to their presumed functions. Excretion cells may represent old digestion cells. The relation between resorption and digestion cells needs further investigation. The epithelium of the posterior part of the midgut consists of two types, transport and secretion cells, which seem to serve mainly for the resorption of water and the secretion of peritrophic membranes, respectively.Peritrophic membranes are secreted by the anterior midgut epithelium mainly in a period between 2 and 4 h after feeding. Chitin or chitin precursors could be localized in vesicles and in the brush border of midgut cells, and in the peritrophic membranes, using colloidal gold labelled with wheat germ agglutinin. Two different textures of chitin-containing microfibrils were found in the peritrophic membranes, either a random or a hexagonal texture. The latter results if the microfibrils polymerize between the basal parts of the microvilli. Irregularities of the hexagonal texture can be correlated with an irregular pattern of the microvilli. In the posterior midgut peritrophic membranes with a random texture, chitin-containing microfibrils are continuously secreted in the form of patches.     

The Development of the Brachiopod Crania (Neocrania) anomala (O. F. Müller)and its Phylogenetic Significance

The freely spawned eggs of Crania go through radial cleavage, embolic gastrulation, and the posteroventral part of the archenteron forms mesoderm through modified enterocoely. The blastopore closes in the posterior end of the larva. The ciliated, lecithotrophic larva has four pairs of coelomic pouches and three pairs of dorsal setal bundles. At metamorphosis, the larva curls ventrally by contraction of a pair of midventral muscles, which are extensions of the first pair of coelomic sacs; the larva attaches by the epithelium just behind the closed blastopore. The brachial valve is secreted by the middle part of the dorsal epithelium and the pedicle valve is secreted by the attachment epithelium. The second pair of coelomic sacs develop small attachment areas at the edge of the dorsal valve and become the lophophore coelom (mesocoel); the third pair of coelomic sacs become the body coelom (metacoel) with the adductor muscles. The posterior position of the closing blastopore is characteristic of deuterostomes. The ventral curving of the settling larva and the formation of both valves from dorsal epithelial areas indicate that the brachiopods have a very short ventral side as opposed to the phoronids. It is concluded that both groups have originated from a creeping ancestor with a straight gut.     

花背蟾蜍蝌蚪变态期角膜发育的研究

作者用光镜和电镜研究了花背蟾蜍蝌蚪变态期角膜的发育。在后肢发育晚期,内、外角膜在中央部位首先愈台,在完全变态期角膜完全愈合,此时角膜上皮细胞增殖,上皮基质变为Bowman’s膜,内、外角膜之间的成纤维细胞和由它分泌的胶原纤维形成角膜基质,内角膜细胞形成单层的角膜内皮,它与角膜基质间的Descemet’s膜最晚形成。     

The female genital system of Ooperipatellus decoratus (Onychophora, Peripatopsidae): an ultrastructural study

The female genital system of the oviparous peripatopsid Ooperipatellus decoratus consists of an ovary, oviducts equipped with receptacula seminis and additional pouches, uteri, and a vagina. It is examined using transmission and scanning electron microscopy. The ovary is made up of paired ovarian tubes united anteriorly and posteriorly and differentiated into a sterile dorsal part and a fertile ventral part with exogenous oocytes. Fertilization presumably occurs in the oviducts once the oocytes pass the receptaculum seminis. Although the receptacula seminis have been reported to occur in juvenile O. decoratus females only, the present study reveals that they are present in adult females as well. Their wall consists of a cuboidal epithelium covered with a thin collagen-muscle layer. The additional pouches are projections of the oviducts facing the receptacula seminis. They are distally closed to the haemocoel by a flattened epithelium and lack external muscle cells. A thin collagen layer is only found proximally. The uteri are characterized by a columnar epithelium with folded cell membranes allowing extension of the uteri, thus facilitating the passage of the large uterine eggs towards the vagina. Another dominating feature of the uteri is a distally increasing secretory production, which probably contributes to chorion development. Cilia occurring along the entire length of the uteri are considered to assist in the transport of eggs towards the vagina.     

Geometric peculiarities of intestinal surface and efficiency of coupling between membrane hydrolysis and transport of nutrients

Kinetics of the membrane hydrolysis of maltose and of the absorption of the released glucose in the isolated loop of rat small intestine has been examined in a wide range of maltose concentrations (25–200 mM) under the conditions of chronic experiments. The processes studied were simulated by means of mathematical models using two approximations of the villous surface of intestinal epithelium: (i) smooth flat surface with adjoining pre-epithelial (“unstirred”) layer and (ii) folded surface with “unstirred” layer between the folds and partly above them. The results of modeling matched well the experimental data in the whole range of maltose concentrations only in the case of the folded surface. A model with this approximation predicts a closer coupling between maltose hydrolysis and absorption of released glucose as well as a lower glucose concentration in the intestinal lumen than in the case of a flat surface. We conclude that in order to evaluate correctly a relative role of various mechanisms of glucose transport across intestinal epithelium under normal conditions, one should take into account the pre-epithelial layer of the small intestine and geometric peculiarities of its epithelial surface.     

Secondary Folding of Olfactory Organ in Young and Adult Sea Trout

The development of the olfactory rosette in parr, smolt and adult sea trout is investigated by means of different fixation and staining techniques. The rosette consists of primary laminae which are folded in four types of secondary laminae. These are here called initial, cuneiform, filiform and fungiform laminae. At least initially secondary folding seems to be an interplay between a blood capillary and the sensory epithelium external to it. The ecological adaptations of folded olfactory organs are discussed.     

Hyolith-type microstructure in a mollusc-like fossil from the Early Cambrian of Yunnan, China

A phosphatized bilaterally symmetrical, spirally coiled conch from the Lower Cambrian phosphates of the Meishucunian Stage of eastern Yunnan, China, is composed of two well-preserved layers. The outer one is built of longitudinal mineralized fibres not strictly parallel to each other, but may anastomose or branch. The fibres form comarginal ribs with transversal grooves between them, corresponding to growth stages. In the ribs, the fibres are arranged closely and are steeply inclined towards the apex, while in the grooves the fibres are subparallel to the conch surface and more loosely packed. Small elliptic and triangular holes are situated within and between the fibres of the outer layer. The inner layer consists of transversal fibres running around the conch. They are almost parallel to each other but partly separated by narrow discontinuous slits. Despite overall morphological similarity of the conch to shells of some early Cambrian molluscs, the observed orthogonal configuration of longitudinally oriented fibrous structures of the outer layer and transversal fibres of the inner layer is found in orthothecid hyoliths, phosphatized microstuctures of which have been described from the Lower Cambrian of the Siberian Platform. The new material from the Yangtze Platform supports a conclusion of a distinctive type of microstructure available in hyoliths different from molluscan microstructures available from the Lower Cambrian.     

Structural organization and epithelial cell types of the intestinal diverticula (protopancreas) of ammocoetes of southern hemisphere lampreys: functional and phylogenetic implications

Larvae of the two southern hemisphere lamprey genera, Mordacia and Geotria, possess one and two intestinal diverticula, respectively, each originating at the oesophageal-intestinal junction. These diverticula comprise an inner layer of simple columnar epithelium composed solely of zymogen and mucous cells, a middle layer consisting mainly of a blood sinus, and an outer serosa layer covered by a simple squamous epithelium (mesothelium). The inner surface is highly folded only in Mordacia. The secretion of mucus probably protects the epithelium from the effects of digestive enzymes secreted by the zymogen cells and/or bile, which enters the diverticulum at its tip. Unlike the situation in southern hemisphere lampreys, the zymogen cells of the larvae of holarctic lampreys are located in the anterior intestine, a condition considered to be primitive. It is thus proposed that intestinal diverticula were developed during the evolution of southern hemisphere lampreys. The relocation of zymogen cells in the diverticula increases the area for these cells, and thus the capacity for the synthesis and secretion of digestive enzymes, particularly in Mordacia where the inner surface is folded.     

Fine structure of the Gene's organ in the camel tick Hyalomma (Hyalomma) dromedarii (Ixodoidea: Ixodidae)

Gene's organ of the camel tick Hyalomma (Hyalomma) dromedarii is located in the anterodorsal region of the body cavity ventrad to the scutum. It consists of a short stalk, dividing posteriorly into 2 pairs of horns and then into tubular glands. In unfed ticks, the epithelial layer of both the stalk and horns is lined internally by 2 cuticular layers; an inner, thin, greatly folded, dense layer surrounds the organ main lumen, and an outer, thick, slightly folded, less dense layer abuts the cell apices. Only the inner cuticular layer extends into the horn posterior region and appears perforated with numerous pore canals and covered with fine, cuticular projections. The horn and tubular glands epithelium is structurally consistent with a secretory function that apparently increases as feeding progresses. During oviposition, the inner cuticular layer unfolds and inflates into a pair of balloonlike structures that evert through the organ external aperture to receive and manipulate each egg as it is laid, coating it with a waxy layer that prevents desiccation. The fine cuticular projections may have a function in gripping the eggs as they leave the vagina. This organ appears to be everted by hydrostatic pressure from the hemolymph and is retracted by muscles.     

Some features of the ultrastructure of the coenecium of Rhabdopleura compacta

Summary The coenecium of Rhabdopleura consists of a series of tubes, some erect and some repent. These tubes are composed of rings, one stacked within another. The rings are smooth on the inside surface and rough outside. Newly laid down rings are thin and smooth on both surfaces, fibrous material is laid down on the external surface during growth in thickness by the cephalic shield of the zooid. The erect tubes remain discrete, but the repent tubes, which are attached to the substratum can become incorporated in a mass of secreted material. The external vertical fibres cross several rings and probably serve to anchor the stack. Besides these fibrils that run for several segments, there are other shorter fibres that run along the length of each cylindrical ring, and are not continuous across the rings. These long and short fibres have features in common with those found in the graptolites.I wish to thank Dr. A. Boyde for scanning electron microscope facilities and for making his expertise so freely available. Dr. A. Stebbing helped me to obtain the specimens. Mrs. E. Bailey and Mr. R. Moss ably provided the technical and photographic assistance     

香鱼消化道及肝脏的形态结构特征

采用解剖及石蜡切片显微技术观察了香鱼消化道及肝脏的组织学结构。香鱼消化道由口咽腔、食道、胃及肠构成。口咽腔大且狭长,其底壁前部有一对粘膜褶,两颌边缘着生宽扁梳状齿,腭骨及舌骨具齿,犁骨无齿;舌由基舌骨突出部分覆盖粘膜构成,舌粘膜上皮为复层扁平上皮,含有较多的杯状细胞和味蕾。食道、胃及肠均由粘膜层、粘膜下层、肌层及外膜构成。食道粘膜层上皮为复层扁平上皮,杯状细胞发达。胃呈V形,由贲门部、胃体部及幽门部组成,胃壁粘膜上皮为单层柱状上皮,贲门部与胃体部的固有层中有胃腺。肠较短,由前、中、后肠构成,肠壁粘膜上皮为单层柱状上皮,其游离面具微绒毛;上皮细胞间有杯状细胞。幽门盲囊有350～400条,其组织学结构与肠相同。肝脏单叶,外被浆膜;肝细胞形态不规则,肝小叶界限不明显。