Periostracal mineralization in the gastrochaenid bivalve Spengleria

We investigated the spikes on the outer shell surface of the endolithic gastrochaenid bivalve genus Spengleria with a view to understand the mechanism by which they form and evaluate their homology with spikes in other heterodont and palaeoheterodont bivalves. We discovered that spike formation varied in mechanism between different parts of the valve. In the posterior region, spikes form within the translucent layer of the periostracum but separated from the calcareous part of the shell. By contrast those spikes in the anterior and ventral region, despite also forming within the translucent periostracal layer, become incorporated into the outer shell layer. Spikes in the posterior area of Spengleria mytiloides form only on the outer surface of the periostracum and are therefore, not encased in periostracal material. Despite differences in construction between these gastrochaenid spikes and those of other heterodont and palaeoheterodont bivalves, all involve calcification of the inner translucent periostracal layer which may indicate a deeper homology.     

How the 'forceps' of Lithophaga aristata (Bivalvia: Mytiloidea) are formed

The calcareous substratum borer, Lithopaga aristata Dillwyn, 1817, secretes posterior incrustations that take the form of interlocking 'forceps'. These are secreted, initially, as asymmetrical ridges by similarly asymmetrical dorsal and ventral glands in the left and right middle folds of the posterior mantle lobes. In the adult, the secretions are more uniformly spread over the posterior shell margins, concealing the juvenile ridges.
Opening and closing of the valves smooths the outer surfaces of the 'forceps' against the burrow wall, which also comes to be lined with calcium carbonate that is reciprocally smoothed, so that they occlude the borehole more effectively. Extension and retraction of the siphons probably smooths the inner surfaces of the 'forceps' which are sharpened by abrasion, one against the other, during valve opening and closing. The pair of inequilateral spikes so produced, project from the burrow aperture, occluding it, but probably, more importantly, distancing aperturally attacking predators from the true posterior shell margin. Interlocking of the 'forceps' and their sharpness may further deter would-be predators.     

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.     

Shell adaptation in achrothelid brachiopods to settlement on a soft substrate

The protegulum of acrothelid inarticulate brachiopods is characterised by slender. elongate, hilaterally arranged spines which are interpreted as adaptations to facilitate settlement on a soft substrate. They provide a mechanism for positioning the gape clear of the substrate and a means of shell eversion to permit recovery from an overturned position. Holoperipheral growth and attainment of a low subconical form by the pedicle valve during later growth stages provide different strategies for solving these problems, and the protegular spines become non-functional and largely worn away. The structures arc illustrated in a species of Orbithele from the Upper Cambrian of Australia.     

Spikey bivalves: intra-periostracal crystal growth in anomalodesmatans

The external shell surfaces of most anomalodesmatan bivalves are studded with small spikes, particularly at the posterior end. We have studied the morphology, mode of growth, and distribution among taxa of these spikes. In this study we found that spikes vary widely in morphology, from acute spikes to flat plaques. Optical and electron microscopy has revealed that the periostraca of Laternula, Myadora, and Thraciopsis consist of an outer dense layer and an inner translucent layer. The dense layer grows at the expense of the inner layer as it progresses toward the shell edge. The spikes begin to grow in the free periostracum, within the translucent periostracal layer, immediately below the dense layer. With growth, they push the dense periostracal layer upward but without penetrating it. Those parts of the spike in contact with this layer cease to grow, which explains the typical conical shape of spikes. When fully grown, spikes reach the base of the translucent layer, becoming incorporated into the outer shell layer. Scanning electron microscopy and electron backscatter diffraction analysis reveal that the spikes of Lyonsia norwegica and Lyonsiella abyssicola are prisms of aragonite composed of twinned crystals, with the c-axis vertical. A survey of the occurrence of spikes within the anomalodesmatans shows that they are present in all but a few families. Elsewhere within the closely related palaeoheterodonts, intra-periostracal calcification is also known in Neotrigonia and unionids, which indicates that this character may be plesiomorphic for these bivalves. The present data do not support the homology of spikes in other bivalve groups (e.g., veneroids) or in the aplacophorans or polyplacophorans.     

Structure of bluetongue virus particles by cryoelectron microscopy.

The structure of the bluetongue virus (BTV) particle, determined by cryoelectron microscopy and image analysis, reveals a well-ordered outer shell which differs markedly from other known Reoviridae. The inner shell is known to have an icosahedral structure with 260 triangular spikes of VP7 trimers arranged on a T = 13,l lattice. The outer shell is seen to consist of 120 globular regions (possibly VP5), which sit neatly on each of the six-membered rings of VP7 trimers. "Sail"-shaped spikes located above 180 of the VP7 trimers form 60 triskelion-type motifs which cover all but 20 of the VP7 trimers. These spikes are possibly the hemagglutinating protein VP2 which contains a virus neutralization epitope. Thus, VP2 and VP5 together form a continuous layer around the inner shell except for holes on the 5-fold axis.     

Large gryphaeid oysters as habitats for numerous sclerobionts: a case study from the northern Red Sea

The shell of a living specimen of the Indo-Pacific gryphaeid giant oyster Hyotissa hyotis was colonized by numerous encrusting, boring, nestling and baffling taxa which show characteristic distribution patterns. On the upper valve, sponge-induced bioerosion predominates. On the lower valve intergrowth of chamid bivalves and thick encrusting associations—consisting mostly of squamariacean and corallinacean red algae, acervulinid foraminifera, and scleractinian corals—provides numerous microhabitats for nestling arcid and mytilid bivalves as well as for encrusting bryozoans and serpulids. Such differences between exposed and cryptic surfaces are typical for many marine hard substrata and result from the long-term stable position of the oyster on the seafloor. The cryptic habitats support a species assemblage of crustose algae and foraminifera that, on exposed surfaces, would occur in much deeper water.     

马传染性贫血病病毒结构模型的建立

通过电子显微镜多种技术的观察和研究,明确了马传染性贫血病病毒的典型形态结构,即呈球形,表面附有顶端带钮状物的纤突,病毒膜为三层(单位膜),病毒基质充填于病毒膜和核芯亮之间,核芯壳呈园锥形,核芯为螺旋结构。在此基础上建立了马传染性贫血病病毒的结构模型。这是国内外首次报道,为深入研究该病毒和其有关病毒的特性奠定了基础,并为该病毒的分类提供了形态学依据。     

Structures of hepatitis B virus cores presenting a model epitope and their complexes with antibodies

The core shell of hepatitis B virus is a potent immune stimulator, giving a strong neutralizing immune response to foreign epitopes inserted at the immunodominant region, located at the tips of spikes on the exterior of the shell. Here, we analyze structures of core shells with a model epitope inserted at two alternative positions in the immunodominant region. Recombinantly expressed core protein assembles into T=3 and T=4 icosahedral shells, and atomic coordinates are available for the T=4 shell. Since the modified protein assembles predominantly into T=3 shells, a quasi-atomic model of the native T=3 shell was made. The spikes in this T=3 structure resemble those in T=4 shells crystallized from expressed protein. However, the spikes in the modified shells exhibit an altered conformation, similar to the DNA containing shells in virions. Both constructs allow full access of antibodies to the foreign epitope, DPAFR from the preS1 region of hepatitis B virus surface antigen. However, one induces a 10-fold weaker immune response when injected into mice. In this construct, the epitope is less constrained by the flanking linker regions and is positioned so that the symmetry of the shell causes pairs of epitopes to come close enough to interfere with one another. In the other construct, the epitope mimics the native epitope conformation and position. The interaction of native core shells with an antibody specific to the immunodominant epitope is compared to the constructs with an antibody against the foreign epitope. Our findings have implications for the design of vaccines based on virus-like particles.     

Localization of the bacterial agent of juvenile oyster disease (Roseovarius crassostreae) within affected eastern oysters (Crassostrea virginica)

The bacterium Roseovarius crassostreae causes seasonal mortalities among commercially produced eastern oysters (Crassostrea virginica) grown in the Northeastern United States. Phylogenetically, the species belongs to a major lineage of marine bacteria (the Roseobacter clade), within which Roseovarius crassostreae is the only known pathogen to be isolated in laboratory culture. The objective of the current study was to determine the location and nature of R. crassostreae interactions with oysters affected by juvenile oyster disease (JOD). Scanning electron microscopy of diseased individuals revealed abundant colonization of the inner shell surfaces by bacteria which were morphologically similar to R. crassostreae. The same types of cells were also observed on and within layers of host-derived conchiolin on the inner valves. Most bacterial cells were alive as determined by the use of a fluorescent viability stain. Further, most were clearly attached at the cell poles, which is consistent with the ability of R. crassostreae to express polar fimbriae. When material from the pallial fluid, soft tissue and inner valve surfaces was cultured, the highest numbers of R. crassostreae were recovered from the inner valves. These samples also contained the greatest abundance of R. crassostreae as a percentage of total colonies. Cloning and sequencing of 16S rRNA genes provided culture-independent evidence of the numerical dominance of R. crassostreae among the bacterial consortia associated with the inner shell surfaces of JOD-affected animals. The ability of R. crassostreae to colonize shell and conchiolin is consistent with the described JOD-pathology and may aid the bacteria in avoiding hemocyte-mediated killing.     

Spreading of mouse fibroblasts on the substrate with multiple spikes

Mouse embryo fibroblasts were cultivated on special substrates with discontinuous surfaces. The substrates were silicon plates with multiple vertical (65-90 microns height) spike-like silicon microcrystals evenly distributed on the plate surfaces. It was shown that the cells were successfully spread and flattened on these substrates. The spread cells formed several discrete attachment zones at the tops and side surfaces of the spikes; these zones were separated from one another by distances considerably greater than the diameter of the unspread cell. At early stages of spreading the unspread cells attached to the tops of single spikes and extended long filopodia attached to the distant spikes. At later stages the lamellae were formed between the filopodia: probably these filopodia served as guidelines for extension of lamellae and progressive cell spreading. These experiments demonstrated that continuity of substrate surface is not a necessary condition for advanced cell spreading.     

Rotavirus spike structure and polypeptide composition.

Negatively stained preparations of rotavirus imaged with a low dose of electrons provide sufficient contrast to reveal surface projections or spikes. The number of spikes found projecting from different particles indicates that not all 60 peripentonal sites are occupied. Treatment at pH 11.2 with 250 mM ammonium hydroxide specifically removes the spikes, yielding smooth double-shelled particles of the same diameter as that of the native virus. Protein analysis confirms that the released spikes are composed of polypeptide VP4 (or its two cleavage products VP5* and VP8*) and that the smooth particle retains the other major outer shell protein VP7. Spikeless particles can be decorated by a monoclonal antibody specific for the major immunodominant neutralizing domain of VP7, implying that removal of the spikes does not denature the VP7 that is retained on the surface of the smooth particle.     

How does flow recruit epibionts onto brachiopod shells? Insights into reciprocal interactions within the symbiotic framework

Computational fluid dynamics simulations were performed to examine the passive recruitment of epibionts onto Devonian spiriferide brachiopod host shells. Because many planktonic larvae and spores are propulsion-inefficient swimmers, we determined the areas most prone to settlement in terms of inertial impaction and direct interception, which are characteristic of higher and lower pressure, respectively. Simulations on a unique specimen of Paraspirifer with a geopetal structure of broken brachidia suggest that the larva of Aulopora on the shell was transported and had settled onto the shell through inertial impaction after the host was dead and overturned on the sea floor. In the case of an ideal life posture, the spiriferide models received higher pressure on the shell surfaces at the forward and rearward stagnation areas and lower pressure along the shell margins and the anterior part of the sulcus, regardless of whether the ventral or the dorsal valve was facing upstream. Both sites seem to be available for epibionts by way of direct interception or inertial impaction. Our results indicate that the initial recruitment of most epibionts is accidental and passive, whereas the directions and patterns of epibiont growth suggest a biological response to ambient conditions.     

Application of freeze-drying intact cells to studies of murine oncornavirus morphogenesis.

Using a method for freeze-drying intact cells, uninfected and murine leukemia virus (MuLV)-infected JLSV9 cell surfaces, as well as murine mammary tumor virus (MuMTV)-infected cell surfaces, were examined by electron microscopy. The 10-nm knobs of MuLV and the 5-nm spikes of MuMTV were clearly revealed on the surfaces of budding viruses and were also found dispersed over the cell surface. The MuLV knobs are randomly arranged on the virus surface, whereas the MuMTV spikes are much more ordered. Because freeze-fractured budding viral envelopes are devoid of intramembranous particles, the observed surface particles do not appear to be merely accentuated intramembranous particles. This technique should permit further analysis of the morphogenesis of viral envelopes without the need for externally applied labels.     

Larval development of the shell and the shell gland inMytilus (Bivalvia)

Summary In the newly hatched larva ofMytilus galloprovincialis the larval shell field almost totally invaginates to form a shell gland. Only very few cells remain outside and they meet at one point. Only these cells secrete the pellicle, which in the beginning is protected by the glycocalyx and the microvilli of the surrounding cells. The invaginated cells do not participate in pellicle secretion. The secretion partly passes through infoldings of the apical cell membrane. Mineralization of each valve begins along the hinge line, continues to the valve margins and includes the valve centre.     

Über Schalenabschliffe beiCardium edule aus der Königsbucht bei List auf Sylt

Two communities of the recent bivalveCardium edule L. were collected in the tidal flats near List on the island of Sylt (Southern North Sea). The cockles were examined for abrasion marks on the surfaces of their valves arising from various causes. One important group of abrasions reveals traction marks restricted to the beak region, caused exclusively by valve movements. Another group is characterized by marks due to shell abrasions on the sea bed. The various types of abrasion marks have been studied in living cockles, both in the sea and in laboratory containers, and the results compared with those obtained on dead ones still connected by the ligament. In the sea, livingCardium edule were studied via Scuba-observations. Traction marks on the umbo support the conclusion that living cockles sometimes rest on their dorsal shell area. Holes in the convex umbo, umbonal facets, originate after death in situ or through shiftings to and fro on the sea bed, and also as a result of chemical or biological processes. The method applied and the criteria of shell abrasions employed are considered acceptable approaches to the interpretation of functional morphology and behaviour in fossil molluscs.     

Earliest ontogeny of Early Palaeozoic Craniiformea: implications for brachiopod phylogeny

Popov, L.E., Bassett, M.G., Holmer, L.E., Skovsted, C.B. & Zuykov, M.A. 2010: Earliest ontogeny of Early Palaeozoic Craniiformea: implications for brachiopod phylogeny. Lethaia, Vol. 43, pp. 323–333. Well preserved specimens of the Early Palaeozoic craniiform brachiopods Orthisocrania and Craniops retain clear evidence of a lecithotrophic larval stage, indicating the loss of planktotrophy early in their phylogeny. The size of the earliest mineralized dorsal shell was <100 μm across, and the well preserved shell structure in these fossil craniiforms allows their earliest ontogeny to be compared directly with that of living Novocrania, in which the first mineralized dorsal shell (metamorphic shell) is secreted only after settlement of the lecithotrophic larvae. Immediately outside this earliest shell (early post‐metamorphic or brephic shell) and in the rest of the dorsal valve the primary layer in both fossil and living craniiforms has characteristic radially arranged laths, which are invariably lacking in the earliest dorsal shell. The ventral valve of the fossil specimens commonly preserves traces of an early attachment scar (cicatrix), which is equal in size to the dorsal metamorphic shell, and the brephic post‐metamorphic ventral valve also has a primary shell with radially arranged laths. However, a primary shell with radial laths is completely lacking in the ventral valve of living Novocrania, indicating that heterochrony may have been involved in the origin of the encrusting mode of life in living craniids; the entire ventral valve of Recent craniids (with the possible exception of Neoancistrocrania) may correspond to the earliest attachment scar of some fossil taxa such as Orthisocrania. It is also probable that the unique absence of an inner mantle lobe as well as the absence of lobate cells in Novocrania could be the result of heterochronic changes. The dorsal valve of both fossil and living craniiforms has a marked outer growth ring, around 500 μm across, marking the transition to the adult, and a significant change in regime of shell secretion. The earliest craniiform attachment is considered to be homologous to the unique attachment structures described recently in polytoechioids (e.g. Antigonambonites) and other members of the strophomenate clade. However, unlike the craniiforms, polytoechioids and strophomenates all have planktotrophic larvae, and planktotrophy is most probably a plesiomorphic character for all Brachiopoda. □Brachiopoda, Craniiformea, Early Palaeozoic, ontogeny, phylogeny.     

Survival and repair of surgical and natural shell damage in the articulate brachiopod Terebratulina retusa (Linnaeus)

Living specimens of Terebratulina retusa from the Firth of Lorn, Scotland, were surgically damaged by drilling 2 mm diameter holes or narrow slits one cm long in the anterior portion of one valve, by bevelling the anterior margin of both valves, or by amputation of the anterior third of one valve. These injuries to the shell and mantle simulated the type of repaired shell damage seen in Paleozoic species, i.e., scalloped, divoted, cleft, and embayed valves. Less than ten percent of the 200 damaged specimens survived until the 25th week after surgery. Specimens of T. retusa showed the ability to repair drill holes, slits, and bevelled anterior shell regions, but not the most severe damage, i.e., amputations of the anterior third of one valve. Shell‐repair was initiated in the fourth week after surgery by the development of a membrane across the wound. The development of caeca in the new shell layer secreted to plug the drill holes became apparent by the eighth week. The punctate pattern was complete in the new, translucent shell material of bevelled and drilled specimens by the 25th week following surgery. Failure of any specimens to survive amputation of the anterior portion of a valve for more than seven weeks after surgery, and the absence of initiation of the repair process, suggests that terebratulids do not have the tolerance for, nor the ability to repair, the severe injuries (embayed valves) which were sustained and mended by extinct strophomenids.     

Is there a link between shell morphology and parasites of zebra mussels?

The shell morphology of zebra mussels, Dreissena polymorpha, was analyzed to determine if alterations in shell shape and asymmetry between valves were related to its infection status, i.e. infected or not by microparasites like ciliates Ophryoglena spp. or intracellular bacteria Rickettsiales-like organisms (RLOs), and by macroparasites like trematodes Phyllodistomum folium and Bucephalus polymorphus. For microparasites, two groups of mussels were observed depending on shell measurements. Mussels with the more concave shells were the most parasitized by ciliates. This could be more a consequence than a cause and we hypothesized that a modification of the water flow through the mantle cavity could promote the infection with a ciliate. There were more RLOs present in the most symmetrical individuals. A potential explanation involved a canalization of the left-right asymmetry as a by-product of the parasite infection. Trematode infections were associated with different responses in valve width. Females infected by P. folium displayed significantly higher symmetry in valve width compared with non-infected congeners, whereas the infection involved an opposite pattern in males. B. polymorphus was also linked to a decrease in valve width asymmetry. This study suggested that a relationship exists between parasitism and shell morphology through the physiological condition of host zebra mussels.     

Morphological and structural adaptations to soft substrates in the Early Jurassic monomyarians Lithiotis and Cochlearites

Lithiotis problematica and Cochlearites loppianus are sessile monomyarian bivalves known from the Early Jurassic of the Tethyan region, where they are found standing vertically in the calcareous mud of lagoonal fades. Their shells are characterized by a long cardinal area, ventrally displaced body space, thick attached valve, and a thin, flat free valve of equal length. A functional ligament is present only in juvenile stages of Cochlearites. To grow straight, the ventral end of the shell had to gape, so in both species it was probably the elasticity of the thin free valve that caused the shell to open. In the adult stage the shell grew only towards the venter with an apparently constant growth rate, while the soft body size remained unchanged. The outer shell is composed of a compact aragonitic layer, while the inner part is filled with loose chalky deposits which are thought to have functioned as supporting the soft body and lifting it upwards. These and other morphologic and structural features, as well as the growth pattern, can be explained as an adaptation of a sessile animal to soft muddy bottoms and rapid sedimentation. Some elongated Crassostrea living in similar environments show remarkable morphologic and structural convergence with Lithiotis and Cochlearites.