STRUCTURE OF THE CONCHIOLIN CASES OF THE PRISMS IN MYTILUS EDULIS LINNE

The prisms in the shell of Mytilus edulis Linné are calcite needles. Their small size and their thin conchiolin cases distinguish them from the prisms of many other species of mollusks. These Mytilus prisms have been studied with the electron microscope. The material consisted of positive replicas of surfaces of the prismatic layer, etched with chelating agents, and of preparations of tubular cases from decalcified prisms which were compared with the conchiolin from decalcified mother-of-pearl of the same species. In the replicas, the cases appear as thin pellicles in the intervals between the prism crystals. Both the prism cases and the nacreous conchiolin, disintegrated by exposure to ultrasonic waves and sedimented on supporting films, appear in the form of tightly meshed, reticulated sheets, described as "tight pelecypod pattern" in former studies on nacreous conchiolin of Mytilus. The results show that in the shell of this species the same conchiolin structure is associated with aragonite in mother-of-pearl and with calcite in the prismatic layer.     

Functional morphology and palaeontological significance of the conchiolin layers in corbulid pelecypods

The Corbulidae, which today are slow, cumbersome, very shallow burrowers, developed special morphological features by which they obtained an outstanding capability to withstand the physical and biological stresses characteristic of their preferred habitat. These features are: an inequivalve, globose shape, thick shells, and conchiolin layers (at least one) embedded within their valves in a unique way. These features enable the corbulids to close their valves tightly during the unfavourable environmental conditions (e.g. low salinity, low oxygen content) which may prevail in the marginal marine regions inhabited by several corbulid species. The conchiolin layers act as a barrier preventing all chemically boring organisms from penetrating into the bivalve shell, or shell dissolution by sea water undersaturated with respect to calcium carbonate. The layered conchiolin weakens the shell mechanically, however, especially during fossilization, when the conchiolin is decomposed. The valve splits apart into two shells so completely different in appearance that they may be attributed to different taxa. The conchiolin layers are therefore of great ecological and palaeontological significance. The nature of these conchiolin layers in Corbula (Varicorbula) gibba (Olivi) is described and illustrated and their functional significance discussed in relation to other living and fossil corbulid species.     

A New Ichnospecies of Gastrochaenolites Leymerie from the Pleistocene Port Morant Formation of Southeast Jamaica and the Taphonomy of Calcareous Linings in Clavate Borings

The ichnospecies Gastrochaenolites pickerilli isp. nov. is based on ten borings found in a shell of the gastropod Strombus gigas Linné from the Pleistocene (Sangamonian) Port Morant Formation of southeast Jamaica. These borings bear morphological similarities to Gastrochaenolites torpedo Kelly and Bromley but differ from all other Gastrochaenolites ispp. in having prominent and numerous calcareous meniscate structures arrayed adjacent to one side of the boring. These menisci are concave towards the center of the boring and are the remnants of calcareous tubes that lined earlier boreholes, that the boring bivalve treated as part of the lithified substrate when relocating. They are thus evidence of the former positions of borings that, unusually, were breached as the bivalve migrated sideways. Although this was a common behavior for Gastrochaenolites-producing bivalves within this substrate, the reason for it occurring is uncertain.     

Characteristics of shell microstructure and growth analysis of the Antarctic bivalve <Emphasis Type="Italic">Laternula elliptica</Emphasis> from Lützow-Holm Bay,Antarctica

Growth performance of the Antarctic bivalve Laternula elliptica was examined both by shell microstructural observation and by applying a fluorescent substance, tetracycline, as a shell growth marker. The shell was composed of two calcareous layers: the thick outer layer was homogeneous or granular in structure and the thin inner layer was nacreous. The architecture of Antarctic L. elliptica was different from that of temperate L. marilina, and the ratio of thickness between the outer and inner layers appeared to be different. The growth rate of the nacreous layer was analyzed to be very low. High correlations were found between the major axis of chondrophore and both shell length and shell dry weight, respectively. It is suggested that the chondrophore is an appropriate growth indicator, and combining the information of growth increments with the fluorescent method may be useful in estimating the bivalve growth performance in the Antarctic sea.     

EVIDENCE FROM THE FOSSIL RECORD OF AN ANTIPREDATORY EXAPTATION: CONCHIOLIN LAYERS IN CORBULID BIVALVES

Conchiolin layers, organic-rich laminae, are characteristic of the shells of corbulid bivalves. The retention of these layers, despite their high metabolic cost, throughout the evolutionary history of Corbulidae has prompted the proposal of several adaptive scenarios to explain the origin and maintenance of these layers. The most widely held hypothesis contends that conchiolin layers are an adaptation for inhibiting drilling by predatory naticid gastropods. However, others suggest that the layers are adaptations to retard shell dissolution in waters undersaturated with calcium carbonate or to increase shell strength in the face of durophagous (shell crushing) predators. In this paper, experiments using recent Corbula (Varicorbula) gibba (Olivi) and observations of corbulids' present natural habitat demonstrate the current utility of conchiolin layers for all three functions: retardation of shell dissolution in waters undersaturated in calcium carbonate, increase of mechanical shell strength, and inhibition of drilling by predatory naticid gastropods. Earlier analyses of the extensive history of naticid predator-corbulid prey interactions suggested that conchiolin layers were an adaptation, a feature that promotes fitness and was built by selection for its current role, for deterring naticid predators. Not only are naticid drillholes widespread in fossil and recent corbulid shells, but an unusually large number of incomplete drillholes terminate unsuccessfully at conchiolin layers. In addition, a phylogenetic analysis of the origin of conchiolin layers and its function to deter naticid predators is consistent with a hypothesis of adaptation for this function. However, this hypothesis is rejected by an examination of fossil Jurassic Corbulomima. These oldest corbulids contained conchiolin layers before the evolution of naticid drilling during the Early Cretaceous. Therefore, conchiolin layers appear to be an exaptation, characters evolved for other usages and later “coopted” for their current role, for defense against drilling predators. The layers may in fact be an adaptation to resist durophagous predation.     

Structure,crystallography, and morphogenesis of the cryptic shell of the terrestrial slug Limax maximus (Mollusca,gastropoda)

The structure and crystallography of the internal shell of the pulmonate gastropod slug Limax maximus were studied at the levels of light and scanning electron microscopy, revealing patterns of shell ontogeny and morphogenesis. The calcified portion of the slightly convex ovoid shell is composed of a single palisade layer of calcitic crystals. Numerous projections, 100 μm in width at the dorsal tip, are found on the dorsal surface of the shell and coincide with local nucleation sites of primordial calcium salt deposition onto the periostracum. With continued calcification these projections coalesce ventrally, forming the single crystalline shell layer. The organic portion of the shell includes the periostracum and an extensive PAS-staining conchiolin. In EDTA-etched preparations, conchiolin appears as a spongy network of fibers throughout the shell. Both horizontal and vertical components of the conchiolin are present, the former of variable thickness and occurring in an intercrystalline manner, the latter always occurring normal to the horizontal set. Macromorphogenic growth is characterized by three distinct temporal stages. Primary growth occurs radially from the umbonal region. Secondary growth is synonymous with shell thickening. Tertiary growth is characterized by both a lateral component, in which the shell extends beyond the primary growth boundaries, and a ventral component, in which the shell continues to grow in thickness. SEM of the ventral shell surface reveals a pattern of growth at the crystalmatrix interface. Proteinaceous fibers of the conchiolin occur unidirectionally in horizontal rows. Zones of incipient calcitic crystallization onto these hypostracal fiber bundles are contrasted by zones of increasing crystallization until the fibrous template (reduced hypostracum) is completely covered by crystals.     

Electron probe analysis of strontium in mussel (Bivalvia,Mytilidae) shells: Feasibility of estimating water temperature

Electron microprobe step-scan analyses across the inner nacreous layer of a sectionedMytilus edulis shell revealed no long-term periodic (e.g., seasonal) variation in the concentration of strontium. Similarly, no significant difference was found between a specimen sampled in February (water temperature = 1.3 °C) and one sampled in August (water temperature = 18.0 °C) with regard to the concentration of strontium within the most recently deposited aragonite. Correlation of the amount of strontium within various nacreous regions of the shells of living or fossil mytilids with water temperatures (present or past) is probably not possible through the use of an electron probe, at least to the extent that strontium variation within the nacre ofMytilus edulis is representative of that in nacreous layers of all mytilids.     

Structural relationship between interlamellar organic sheets and nacreous tablets in gastropods and the cephalopodNautilus

The nacreous tablets in gastropods and the cephalopodNautilus are composed of three calcareous layers: a principal, thick, finely granular layer and two thin, coarse-granular layers, one covering the upper surface of the principal layer and another the lower surface of this layer. The granules on the surface layers inNautilus differ from those in gastropods by their much more elongated shape and larger size. The central portion of the nacreous tablet of gastropods andNautilus is more or less elevated forming the central elevation. The granules on this portion usually are larger, irregularly shaped and more crowded than on the main, peripheral portion of the tablet. The untreated, dry interlamellar organic sheets on upper surfaces of immature nacreous tablets are uncalcified and elastic. Narrow thicker parts of the sheet, the trabeculae, Surround large intertrabecular spaces where the sheet is thin. In places it can be observed that each calcareous granula on the surface layer of the nacreous tablet is situated within the intertrabecular space. The size, shape and distribution of the intertrabecular Spaces correspond those of the surface granules. No mineral bridges were observed between the consecutive nacreous tablets.      

Crystalline structure,orientation and nucleation of the nacreous tablets in the cephalopod <Emphasis Type="Italic">Nautilus</Emphasis>

The nacreous tablets in the Nautilus shell have similar crystalline structure as the tablets in the gastropod Gibbula shell. Etching with Mutvei’s solution reveals that each tablet is composed of vertical crystalline columns that are structurally similar to the acicular crystallites in the outer spherulitic-prismatic layer of the shell wall. The columns are attached to each other to form numerous vertical crystalline lamellae, oriented parallel to the longitudinal axis of the tablet. It is still unknown whether or not the orientation of the vertical lamellae corresponds to that of the crystallographic a- or b-axis. The orientation of the crystalline lamellae in the adjacent tablets is parallel in some nacreous laminae, but random in other laminae. Similar large variation was found in the nacreous tablets of the gastropod and bivalve shells. The nucleation sites of the nacreous tablets are predominantly situated on the peripheral portion of the upper surface of the preceding tablet, both in the shell wall and septa. The “aragonite-nucleating proteins” in the central portion of the crystal imprints of the organic interlamellar sheets, described by several writers, have therefore a negative correlation with the nucleation sites of the nacreous tablets.     

Studies on shell formation. VIII. Electron microscopy of crystal growth of the nacreous layer of the oyster Crassostrea virginica

Electron microscope observations have been made by means of the replica method on growth processes of calcite crystals of the nacreous layer of the shell of the oyster, Crassostrea virginica. Layer formation is initiated by the secretion of a conchiolin matrix and the deposition of rounded crystal seeds on or in this material. In some areas crystal seeds are elongate and within a given area show a similar orientation, probably due to slower deposition. The seeds appear to increase in size by dendritic growth, and smaller seeds become incorporated into larger ones which come into contact to form a single layer. With further growth, crystals overlap, forming a step-like arrangement. The direction of growth is frequently different in neighboring regions. Crystal seeds deposited on crystal surfaces are usually elongate and oriented. Well developed crystals have a tabular idiomorphic form and are parallel in their growth. Rounded and irregular crystals were also observed. The crystals show reticular structure with units of the order of 100 A and striations corresponding with the rhombohedral axes of the crystals. The role of the mantle is discussed in relation to the growth patterns of crystals and shell structure.     

Bioerosion of gastropod shells: with emphasis on effects of coralline algal cover and shell microstructure

Organisms boring into fifty nine species of gastropod shells on reefs around Guam were the bryozoan Penetrantia clionoides; the acrothoracian barnacles Cryptophialus coronorphorus, Cryptophialus zulloi and Lithoglyptis mitis; the foraminifer Cymbaloporella tabellaeformis, the polydorid Polydora sp. and seven species of clionid sponge. Evidence that crustose coralline algae interfere with settlement of larvae of acrothoracian barnacles, clionid sponges, and boring polychaetes came from two sources: (1) low intensity of boring in limpet shells, a potentially penetrable substrate that remains largely free of borings by virtue of becoming fully covered with coralline algae at a young age and (2) the extremely low levels of boring in the algal ridge, a massive area of carbonate almost entirely covered by a layer of living crustose corallines. There was a strong negative correlation between microstructural hardness and infestation by acrothoracian barnacles and no correlation in the case of the other borers. It is suggested that this points to a mechanical rather than a chemical method of boring by the barnacles. The periostracum, a layer of organic material reputedly a natural inhibitor of boring organisms, was bored by acrothoracican barnacles and by the bryozoan. The intensity of acrothoracican borings is shown to have no correlation with the length of the gastropod shell.     

An insect boring in an Early Cretaceous wood from Bornholm,Denmark

Abstract

An insect boring of unique shape is described from a lignitic layer within the Early Cretaceous (Berriasian) Skyttegård Member of the Rabekke Formation on Bornholm. Morphologically it cannot be compared to any modern or fossil wood borings, although some structures are reminiscent of Scolytidae, Platypodidae and Lymexylonidae. Most probably, however, the tracemaker was a female fungus-farming beetle, thus producing an agrichnion.     

Skeletal chemistry of Nautilus and its taxonomic significance

The strontium (Sr) and magnesium (Mg) chemistry of the shell wall and septum as well as the spherulitic-prismatic and nacreous layers of these structures was determined for Nautilus species: N. belauensis, N. macromphalus, N. pompilius and N. scrobiculatus. Each species of Nautilus exhibits greater variability and higher concentrations of Mg in juvenile portions of the shell than in more mature portions of the shell. This decrease in the variability and amount of Mg in the aragonite lattice suggests a physiochemical system which becomes more efficient with time relative to carbonate production. Statistically significant differences in the Sr and Mg content of spherulitic-prismatic and nacreous layers of the shell and septum indicate that these layers were formed from extracellular fluids of different compositions. Concentrations of Sr and Mg in aragonite of the shell wall are characteristic for each species and sufficiently invariant within species to allow species of Nautilus to be distinguished statistically on the basis of either the Sr or Mg content of the shell wall or the Mg content of septa.     

Structure and composition of the septal nacreous layer of Nautilus macromphalus L. (Mollusca, Cephalopoda)

The nacreous layer of Mollusca is the best-known aragonitic structure and is the usual model for biomineralization. However, data are based on less than 10 species. In situ observations of the septal nacreous layer of the cephalopod Nautilus shell has revealed that the tablets are composed of acicular laths. These laths are composed of round nanograins surrounded by an organic sheet. No hole has been observed in the decalcified interlamellar membranes. A set of combined analytical data shows that the organic matrices extracted from the nacreous layer are glycoproteins. In both soluble and insoluble matrices, S amino acids are rare and the soluble organic matrices have a higher sulfated sugar content than the insoluble matrices. It is possible that the observed differences in the structure and composition of the nacreous layers of the outer wall and septa of the Nautilus shell have a dual origin: evolution and functional adaptation. However, we have no appropriate data as yet to answer this question.     

The Fossil Record of Shell Boring by Snails

The predatory boring habit common to many recent snails probablyarose first in the Polinicinae (Naticacea) during Upper Cretaceous(Cenomanian) times (100 million years B.P.) . In the fossilrecord the frequency of bored shells increasesgreatly in rocksof latest Cretaceous age and becomes more widespread duringearly Tertiary times coincident with the major diversificationof the primary groups of boring snails. The borings in these Cretaceous and Tertiary shells show thesame characteristics of preference of penetration in one pelecypodvalve rather than the other or in position of the boring siteon the shell that are found in recent shell assemblages. Borings in Paleozoic brachiopod shells (230–550 millionyears old) that have previously been attributed to gastropodpredation are herein attributed to other but unknown boringorganisms. In part these borings are not accepted as evidence of Paleozoicgastropod predation because it necessitates: (1) Postulationof the separate development of a boring habit ith its concomitantdevelopment of an accessory boring organ in a groupwhose descendantsare all herbivores, and (2) The development of such a habithundreds of millions of years before the appearance of any relativesof present day borers.     

Ultrastructural Characteristics of the Nacre in Some Gastropods

The nacreous layer in Gibbula, Calliostoma, Trochus and Haliotis is described on the basis of scanning electron microscopic studies. The central part of each nacreous tablet contains a significant amount of calcified organic matrix which is insoluble in a chromium sulphate and a 25% glutaraldehyde solution. In most cases, the tablet is subdivided by radial vertical organic membranes into a varying number (2 to 50) of crystalline sectors. These sectors represent polysynthetically twinned crystal individuals which form cyclic or interpenetrant twins. The nacreous tablets in gastropods are compared with those in bivalves, and with the non-biogenic aragonite. The mechanical properties of the nacre, and the effects of the interlamellar conchiolin membranes upon the nucleation of the tablets, are discussed.     

Aragonitic dendritic prismatic shell microstructure in Thracia (Bivalvia,Anomalodesmata)

The shells of most anomalodesmatan bivalves are composed of an outer aragonitic layer of either granular or columnar prismatic microstructure, and an inner layer of nacre. The Thraciidae is one of the few anomalodesmatan families whose members lack nacreous layers. In particular, shells of members of the genus Thracia are exceptional in their possession of a very distinctive but previously unreported microstructure, which we term herein “dendritic prisms.” Dendritic prisms consist of slender fibers of aragonite which radiate perpendicular to, and which stack along, the axis of the prism. Here we used scanning and transmission electron microscopical investigation of the periostracum, mantle, and shells of three species of Thracia to reconstruct the mode of shell calcification and to unravel the crystallography of the dendritic units. The periostracum is composed of an outer dark layer and an inner translucent layer. During the free periostracum phase the dark layer grows at the expense of the translucent layer, but at the position of the shell edge, the translucent layer mineralizes with the units typical of the dendritic prismatic layer. Within each unit, the c‐axis is oriented along the prismatic axis, whereas the a‐axis of aragonite runs parallel to the long axis of the fibers. The six‐rayed alignment of the latter implies that prisms are formed by {110} polycyclically twinned crystals. We conclude that, despite its distinctive appearance, the dendritic prismatic layer of the shell of Thracia spp. is homologous to the outer granular prismatic or prismatic layer of other anomalodesmatans, while the nacreous layer present in most anomalodesmatans has been suppressed.     

Studies on Shell Formation : VIII. Electron Microscopy of Crystal Growth of the Nacreous Layer of the Oyster Crassostrea virginica

Electron microscope observations have been made by means of the replica method on growth processes of calcite crystals of the nacreous layer of the shell of the oyster, Crassostrea virginica. Layer formation is initiated by the secretion of a conchiolin matrix and the deposition of rounded crystal seeds on or in this material. In some areas crystal seeds are elongate and within a given area show a similar orientation, probably due to slower deposition. The seeds appear to increase in size by dendritic growth, and smaller seeds become incorporated into larger ones which come into contact to form a single layer. With further growth, crystals overlap, forming a step-like arrangement. The direction of growth is frequently different in neighboring regions. Crystal seeds deposited on crystal surfaces are usually elongate and oriented. Well developed crystals have a tabular idiomorphic form and are parallel in their growth. Rounded and irregular crystals were also observed. The crystals show reticular structure with units of the order of 100 A and striations corresponding with the rhombohedral axes of the crystals. The role of the mantle is discussed in relation to the growth patterns of crystals and shell structure.     

Identification of chitin in the prismatic layer of the shell and a chitin synthase gene from the Japanese pearl oyster, Pinctada fucata

The shell of the Japanese pearl oyster, Pinctada fucata, consists of two layers, the prismatic layer on the outside and the nacreous layer on the inside, both of which comprise calcium carbonate and organic matrices. Previous studies indicate that the nacreous organic matrix of the central layer of the framework surrounding the aragonite tablet is beta-chitin, but it remains unknown whether organic matrices in the prismatic layer contain chitin or not. In the present study, we identified chitin in the prismatic layer of the Japanese pearl oyster, Pinctada fucata, with a combination of Calcofluor White staining with IR and NMR spectral analyses. Furthermore, we cloned a cDNA encoding chitin synthase (PfCHS1) that produces chitin, contributing to the formation of the framework for calcification in the shell.