Skeletal formation in the modern but ultraconservative chaetetid spongeSpirastrella (Acanthochaetetes) wellsi (demospongiae,porifera)

Summary The modern hadromerid coralline spongeSpirastrella (Acanthochaetetes) wellsi exhibits a unique secondary high-Mg calcite (>19 mol % MgCO₃) basal skeleton. The basal skeleton is constructed of bundles of elongated crystals more or less tangentially orientated. The initial formation of these crystals is controlled by soluble highly acidic aspartic and glutamic-rich (40%) macromolecules. The skeletal mineralization occurs in four different loci: in the top of the calicles, at the tabulae, on collagenous anchor fibres, and within closed spaces between the tabulae. The clicle walls are formed on the uppermost top of the basal skeleton as a continuous process. Based on long term stainings with Ca²⁺-chelating fluorochroms (calcein, chlorotetracyclines) the growth rate of this sponge is extremely low with ca. 50–100μm/a. The skeletal formation takes places outside the sponge, within a narrow zone (300–500 nm) between the basopinacoderm and the mature basal skeleton. The sponge produces thread-like folded templates (‘spaghetti fibres’) of 0,5–2 μm size, the shape controlling insoluble organic matrix. These templates become mineralized in a first step as MgCO₃, then are stretched. A soluble organic matrix is also secreted, and remains are included inside the mineralized skeleton. This organic matrix consists of in a complex mixture containing small very acidic proteins (5, 13, 31 KD; 40% Asp and Glu and therefore most probably Ca²⁺-binding) and high molecular weight glycoproteins among several other organic compounds. The mature crystals are high-Mg calcites. During calcification large cells with large reserve granules (LCG) are always present in a tight connection with the basopinacoderm. These cells form also the collagenous anchor fibres. Primary tabulae are formed by a non-collagenous organic sheet. Calcification happens only when LCG cells are enriched on the organic sheet. Randomly oriented high-Mg calcite crystals are growing on the collagenous anchor fibres. The same type of the mineralization is observed within the spaces of the tabulae. This particular case of mineralization is controlled by decaying sponge tissue (ammonification). The δ¹³C values are in equilibrium with the ambient sea water and vary between +3.2 and +2.8 ‰. The mode of mineralization of the basal skeleton can be described as biologically induced resp. matrix mediated.     

Spicule formation in the calcareous sponge Sycon ciliatum

Summary The spicule primordium is formed in an intercellular cavity within a group of sclerocytes. This cavity contains organic material which ensheaths the growing spicule but does not appear to determine the nature of the mineral morph (magnesian calcite) or the crystallographic orientation of the spicule. The tip of each growing spicule ray is seated in a dense cup in the cytoplasm of the sclerocyte concerned. Both ends of monaxons are initially inserted each into a dense cup. As rays elongate the sclerocyte membrane around the tip becomes invaginated and forms a system of converging spaces that possibly indicate high secretory activity in that region. Spicule growth involves the displacement and expansion of the organic sheath by the enlarging spicule. Fully formed spicules which are exposed to the mesohyl become surrounded by collagen fibrils. However, these fibrils are in no way concerned with the process of mineral deposition and are never found within the spicule calcite.     

Calcite crystal orientation patterns in the bilayers of laminated shells of benthic rotaliid foraminifera

Shells of calcifying foraminifera play a major role in marine biogeochemical cycles; fossil shells form important archives for paleoenvironment reconstruction. Despite their importance in many Earth science disciplines, there is still little consensus on foraminiferal shell mineralization. Geochemical, biochemical, and physiological studies showed that foraminiferal shell formation might take place through various and diverse mineralization mechanisms.In this study, we contribute to benthic foraminiferal shell calcification through deciphering crystallite organization within the shells. We base our conclusions on results gained from electron backscattered diffraction (EBSD) measurements and describe microstructure/texture characteristics within the laminated shell walls of the benthic, symbiontic foraminifera: Ammonia tepida, Amphistegina lobifera, Amphistegina lessonii. We highlight crystallite assembly patterns obtained on differently oriented cuts and discuss crystallite sizes, morphologies, interlinkages, orientations, and co-orientation strengths.We show that: (i) crystals within benthic foraminiferal shells are mesocrystals, (ii) have dendritic-fractal morphologies and (iii) interdigitate strongly. Based on crystal size, we (iv) differentiate between the two layers that comprise the shells and demonstrate that (v) crystals in the septa have different assemblies relative to those in the shell walls. We highlight that (vi) at junctions of different shell elements the axis of crystal orientation jumps abruptly such that their assembly in EBSD maps has a bimodal distribution. We prove (vii) extensive twin-formation within foraminiferal calcite; we demonstrate (viii) the presence of two twin modes: 60°/[0 0 1] and 77°/~[6 –6 1] and visualize their distributions within the shells.In a broader perspective, we draw conclusions on processes that lead to the observed microstructure/texture patterns.     

Coccolith crystals ofPleurochrysis carterae: Crystallographic faces, organization, and development

Summary The crystallographic and morphological configuration of the mineral ring associated with the coccoliths ofPleurochrysis carterae was determined by transmission electron microscopy and electron diffraction. Mature Pleurochrysis coccoliths consist of an oval organic base plate, a distal rim of interlocking calcite crystals, and a narrow ribbon of organic material which tethers the mineral ring to the base plate. Crystals of two distinct forms (R and V units) alternate about the rim in a quasi regular manner; their crystallographicc-axes are aligned parallel to and inclined about 63° to the coccolith plane, respectively. The mineral ring has four platelike elements: the distal-shield and outer-tube elements which form the V unit, and the proximal-shield and inner-tube elements which form the R units. The platy surfaces of both tube elements correspond to the common (10 4) rhombohedral faces of calcite, and the plates of the proximal-shield element are prismatic (2 0) faces. The plates of the distal-shield element are rather curved and their orientation does not correspond to a favorable calcite face; however, for convenience they are described as approximately ( 108) faces, faces which rarely, if ever, develop in inorganic sources of calcite. During coccolith development the earliest habits observed for both V and R units correspond to rectangular parallelepipeds. Outgrowth from the initial V unit begins by expansion of (10 4) faces which form the platy surfaces of the outer-tube element. Throughout this period of development the mineral ring is flexible, at least in an isolated state. Subsequent outgrowth of the inner-tube and proximal-shield elements from the initial R unit produces a rigid interlocking ring. The unusual ( 108) faces of the distal-shield element develop after the crystals are locked in place. Organic structures in intimate association with the mineral phase during its nucleation and growth include the coccolith ribbon, the calcium-polyanion particles, and the membrane of the coccolith vesicle. These structures are described in reference to their putative functions in regulating the development of V and R units.Abbreviation PS polysaccharide     

Analyses archéométriques dans la grotte ornée Mayenne-Sciences (Thorigné-en-Charnie, Mayenne)

Since 1998, a multidisciplinary team works on the study of the representations in the cave Mayenne-Sciences (Thorigné-en-Charnie, Mayenne). Particularly, there are trials for U-TH (TIMH) datations of the speleothems recovering the drawings and fossils of the bats; this would help to know at what date the decorated cave was closed. The studies already allowed to know better the black drawings, executed with a wooden charcoal crayon. Thus, it was possible to make removals which gave two dates from the Gravettian phase; this feeds again the discussion about the chronocultural position of this cave, on of the most septentrional caves we know.     

Construction and nanomechanical properties of the exoskeleton of the barnacle, Amphibalanus reticulatus

Barnacles are some of the major inhabitants of intertidal zones and have calcite-based exoskeleton to anchor and armor their tissues. Structural characterization studies of the specie Ambhibalanus reticulatus were performed to understand the construction of the exoskeleton which forms a light-weight yet stiff structure. The parietal shell is constructed of six compartments to yield a truncated cone geometry, which is neatly fixed onto the basal shell that attaches the organism to the substrate surface. The connections among the different compartments happen through sutured edges and also have chemical interlocking to make the junctions impermeable. Also, the shell parts are furnished with hollow channels reducing the overall mass of the construction. The structure and functions of different parts of the exoskeleton are identified and outlined. Finally, the mechanical properties such as modulus, hardness and fracture toughness of the exoskeleton obtained by indentation techniques are discussed.     

Polykristalliner Calcit bei Seeigeln (Echinodermata,Echinoidea)

Zusammenfassung Es wurde erstmals für Echinodermen primär polykristalliner Calcit nachgewiesen, und zwar im Cortex der Primärstacheln der Cidaridae, dem sekundären Zahnskelet von Clypeaster und in den akzessorischen Kalkstrukturen, die im Kauabschnitt die Furche der Diadematiden-Zähne ausfüllen. Es gibt bei anderen Seeigelfamilien keine Bildungen, die dem Cidariden-Cortex oder den akzessorischen Kalkstrukturen der Diadematiden homolog sind. Das polykristalline sekundäre Zahnskelet von Clypeaster ist dagegen dem monokristallinen sekundären Zahnskelet anderer Seeigel homolog.Der Mg-Gehalt des Calcits liegt in den feinkristallinen Zonen (mit Ausnahme des Cortex) im allgemeinen höher; die höchsten Werte finden sich in den Steinteilen der Zähne, gleichgültig ob das sekundäre Zahnskelet mono- oder polykristallin ist.Polykristalline Teile sind im allgemeinen härter als monokristalline Teile. Die Steinteile der Seeigelzähne sind die härtesten Skeletteile von Echinodermen überhaupt; ihre VickersHärte übertrifft weit diejenige von solidem Calcit. Im Steinteil ist das feinkristalline Gefüge von Calcit eng mit organischer Matrix verbunden, und es wird vermutet, daß darauf die besonders hohe Härte der Steinteile beruht.

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Polycristalline calcite in sea urchins

Summary For the first time primary polycrystalline calcite in Echinoderms is shown in the cortex of primary spines of Cidaridae, in the secondary tooth skeleton of Clypeaster and in the accessory calcareous structures filling the crevice fold in the chewing areas of Diadematoidae teeth. Other Echinoid families lack formations homologous to the cortex of Cidaridae and accessory calcareous structures of Diadematoidae. On the other hand the polycrystalline secondary tooth skeleton of Clypeaster is homologous to the monocrystalline one of the other sea urchins.With the exception of cortex the Mg-content in calcite—analyzed by microprobe and X-ray powder method—is generally greater in macrocrystalline parts. The highest Mg-contents are found in the stone parts of teeth irrespective of whether the secondary tooth skeleton is monocrystalline or polycrystalline.Polycrystalline parts are usually harder than monocrystalline ones. The stone parts of Echinoid teeth are the hardest skeleton parts of Echinoderms on the whole; their hardness is much greater than that of solid calcite. It is supposed that the strong interlacing of the microcrystalline calcite and organic matter causes the enormous hardness of the stone part.

Mit Unterstützung durch die Deutsche Forschungsgemeinschaft.     

Spatial distribution of calcite and amorphous calcium carbonate in the cuticle of the terrestrial crustaceans Porcellio scaber and Armadillidium vulgare

The crustacean cuticle is an interesting model to study the properties of mineralized bio-composites. The cuticle consists of an organic matrix composed of chitin–protein fibres associated with various amounts of crystalline and amorphous calcium carbonate. It is thought that in isopods the relative amounts of these mineral polymorphs depend on its function and the habitat of the animal. In addition to the composition, the distribution of the various components should affect the properties of the cuticle. However, the spatial distribution of calcium carbonate polymorphs within the crustacean cuticle is unknown. Therefore, we analyzed the mineralized cuticles of the terrestrial isopods Armadillidium vulgare and Porcellio scaber using scanning electron-microscopy, electron probe microanalysis and confocal μ-Raman spectroscopic imaging. We show for the first time that the mineral phases are arranged in distinct layers. Calcite is restricted to the outer layer of the cuticle that corresponds to the exocuticle. Amorphous calcium carbonate is located within the endocuticle that lies below the exocuticle. Within both layers mineral is arranged in rows of granules with diameters of about 20 nm. The results suggest functional implications of mineral distribution that accord to the moulting and escape behaviour of the animals.