The Environment Recording Unit in coral skeletons – a synthesis of structural and chemical evidences for a biochemically driven, stepping-growth process in fibres

International audienceThis paper gathers a series of structural and biochemical in situ characterizations carried out to improve our knowledge of the fine scale growth patterns of fibres in coral skeletons. The resulting data show a clear correspondence between the mineral subunits of fibres and the spatial distribution of organic macromolecules. New observations using atomic force microscope confirm the close relationship between mineral and organic phases at the nanometre scale. Synthesis of these data results in a significant change in our concept of the mineralization process in coral skeletons. In contrast to the usual view of an aggregate of purely mineral units independently growing by simple chemical precipitation, coral fibres appear to be fully controlled structures. Their growth process is based on cyclic secretion of mineralizing compounds by the polyp basal ectoderm. These biochemical components of the coral fibres, in which sulfated acidic proteoglycans probably play a major role, are repeatedly produced (proteoglycans are those glycoproteins whose carbohydrate moieties consist of long unbranched chains of sulfated amino sugars). This results in a stepping growth mode of fibres and a layered global organization of coral skeletons. Therefore, in contrast to the widely accepted geochemical interpretation, we propose a fibre growth model that places coral skeletons among the typical ''matrix mediated'' structures. The crystal-like fibres are built by superimposition of few micron-thick growth layers. A biomineralization cycle starts by the secretion of a mineralizing matrix and the final step is the crystallization phase, during which mineral material grows onto the organic framework. Thus, each growth layer is the actual Environment Recording Unit. From a practical standpoint, these results may contribute to develop a new high resolution approach of the environment recording by coral skeletons

Oxygen isotope equilibrium in brachiopod shell fibres in the context of biological control

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The environment recording unit in coral skeletons: structural and chemical evidences of a biochemically driven stepping-growth process in coral fibres

International audienceTo improve our understanding of the environment recording by scleractinian corals, a detailed study of the skeleton microstructure has been carried out. A series of physico-chemical in situ characterizations was made, an approach that provides us with structural and biochemical information at the micrometric and nanometric scales. Gathering of these data results in a significant change in our concept of the growth of coral skeletons. In contrast to the usual view of an aggregate of purely mineral units (the coral fibres) independently growing by a simple chemical precipitation, coral skeletons appear to be biochemically controlled structures. Both structural and biochemical data reveal the micron-scaled stepping growth-mode of fibres, and its global coordination. In this process, sulfated acidic proteoglycans probably play a major role, due to their ability to create polymeric frameworks. Atomic force microscopy confirms the close relationship of organic and mineral phases at the nanometric scale. A new microstructural model of coral skeleton formation is proposed, that places coral skeletons among the typical "matrix mediated structures". From a practical standpoint, these results may contribute to develop a new high resolution approach in the study of paleoenvironments

Subdaily and hourly growth patterns within the shell of the chilean gastropod <em>Concholepas concholepas</em> : new perspectives for high-resolution sclerochronological studies

International audienceUnlike many mollusc species, shells of Concholepas concholepas [Bruguière, 1789] do not show obvious annual morphological marks, neither at the macroscopic or the microscopic scales. The coarsely reticulated structure of the outer shell of this species is produced by the intersection of protuberant lamellae and radial ribs stemming from the apex. These lamellae do not seem to have a particular chronological significance, as they can be formed at rather constant intervals during the first year of growth, and at a diminishing frequency during the successive following years. Therefore, the counting of these concentric lamellae is of little help to determine the age of the shells, and the mechanism responsible for their formation still remain to be understood. At a microscopic level, thin sections of a transversal cut of a shell clearly exhibit that it is made up of two principal components: an external layer formed by prismatic crystals of calcite whose thickness increases from the apex toward the paleal edge and an internal aragonitic layer organized in a crossed lamellar structure. Under appropriate amplification and with a particular combination of transmitted and reflected light, the external layer shows growth structures with an alternating brownish and lighter coloured banding. These structures whose differences in colour seem to depend on their organic matter content typically measure between 20 and 150 µm in thickness. On another hand, scanning electron microscopy analyses reveal that the smallest growth structure consists in 2 µm-thick increments. Even though it has not been possible to directly “see” grouping of these bio-mineralisation units under SEM, it could be deduced that the brownish and light-grey bands are thus formed by a varying number of 2 µm-thick increments. Repeated staining experiments realized with fluorochromes (calcein) on living shells provided a precise chronological framework for the understanding of the growth modality of this species. The calcein marks allowed us to establish that the alternating bands correspond to day and night growth increments. The relative thickness of the bands depends on the number of elementary biomineralisation units or, in other terms, on the frequency of formation of these units. Within one of the conducted experiments which lasted a month and a half, it could be demonstrated that the rhythm at which the organism formed these 2 µm-thick units varied between half-an-hour to two hours. Understanding of the growth modality and the possibility to measure sub-daily increments within the shell of C. concholepas open new perspectives for a series of studies on the relationships between environmental conditions and shell growth on this species. It thus becomes possible to work now on variations of chemical composition of the calcitic increments (calibration of geochemical proxies for environmental conditions) at a time scale of the order of a few hours. This study was financed and conducted in the frame of the French PNEDC CONCHAS Project and also benefited from the support of the EU-project CENSOR (Climate variability and El Nino Southern Oscillation: Impacts for natural resources and management, contract 511071). It is CENSOR publication 0093. Contact Information: N. Guzmán, UR Paléotropique, Centre IRD-Ile de France, 32 avenue Henri Varagnat, 93143 Bondy Cedex, France. Phone: 00 331 48 02 55 88, Fax: 00 331 48 02 55 54, Email: [email protected]

Un Spongiaire Sphinctozoaire colonial apparenté aux constructeurs de récifs triasiques survivant dans le bathyal de Nouvelle-Calédonie

Un second représentant actuel des Sphinctozoaires, importants constructeurs de récifs au Permo-Trias, a été découvert dans la zone bathyale de la NouvelleCalédonie. Contrairement au survivant déjà connu, #Valecetia crypta$, il a conservé le mode de croissance colonial et les capacités constructrices de ses analogues fossiles. Sa croissance est bien plus lente que celle des coraux récifaux actuels. La base d'une construction de 10 cm d'épaisseur a été datée de 700 ans. (Résumé d'auteur

Structural, mineralogical, and biochemical diversity in the lower part of the pearl layer of cultivated seawater pearls from Polynesia

A series of Polynesian pearls has been investigated with particular attention to the structural and compositional patterns of the early developmental stages of the pearl layer. These initial steps in pearl formation bear witness of the metabolic changes that have occurred during the pearl-sac formation. The resulting structurally and biochemically complex structures have been investigated using a variety of techniques that provide us with information concerning both mineral phases and the organic components. Results are discussed with respect to our understanding of the biomineralization mechanisms, as well as for the grafting process

Mitochondrial and nuclear genes suggest that stony corals are monophyletic but most families of stony corals are not (Order Scleractinia, Class Anthozoa, Phylum Cnidaria)

Modern hard corals (Class Hexacorallia; Order Scleractinia) are widely studied because of their fundamental role in reef building and their superb fossil record extending back to the Triassic. Nevertheless, interpretations of their evolutionary relationships have been in flux for over a decade. Recent analyses undermine the legitimacy of traditional suborders, families and genera, and suggest that a non-skeletal sister clade (Order Corallimorpharia) might be imbedded within the stony corals. However, these studies either sampled a relatively limited array of taxa or assembled trees from heterogeneous data sets. Here we provide a more comprehensive analysis of Scleractinia (127 species, 75 genera, 17 families) and various outgroups, based on two mitochondrial genes (cytochrome oxidase I, cytochrome b), with analyses of nuclear genes (ßtubulin, ribosomal DNA) of a subset of taxa to test unexpected relationships. Eleven of 16 families were found to be polyphyletic. Strikingly, over one third of all families as conventionally defined contain representatives from the highly divergent "robust" and "complex" clades. However, the recent suggestion that corallimorpharians are true corals that have lost their skeletons was not upheld. Relationships were supported not only by mitochondrial and nuclear genes, but also often by morphological characters which had been ignored or never noted previously. The concordance of molecular characters and more carefully examined morphological characters suggests a future of greater taxonomic stability, as well as the potential to trace the evolutionary history of this ecologically important group using fossils

Evidence of a scheduled end for prism growth in the shell of Pinctada margaritifera : closure of the calcite biomineralization area by a specific organic membrane

The shell of the pearl oyster Pinctada margaritifera is made up of two layers: an outer layer of calcite prisms and an inner layer of aragonite tablets. Recent studies have shown that the calcite layer develops in a series of steps. We found that the end of prism growth and the start of aragonite deposition are also complex processes. Contrary to the common belief that prism growth is interrupted by the expansion of the aragonite layer, we found that a specific membrane covers the calcite surface before aragonite deposition starts. The earliest aragonite depositions occur as granular spots located only on the surfaces covered by this organic membrane This membrane appears to be the final stage of the calcite biomineralization cycle. This new understanding of calcite development has implications for shell biomineralization research and the production of pearls

Limpet Shells from the Aterian Level 8 of El Harhoura 2 Cave (Témara, Morocco): Preservation State of Crossed-Foliated Layers

International audienceThe exploitation of mollusks by the first anatomically modern humans is a central question for archaeologists. This paper focuses on level 8 (dated around * 100 ka BP) of El Har-houra 2 Cave, located along the coastline in the Rabat-Témara region (Morocco). The large quantity of Patella sp. shells found in this level highlights questions regarding their origin and preservation. This study presents an estimation of the preservation status of these shells. We focus here on the diagenetic evolution of both the microstructural patterns and organic components of crossed-foliated shell layers, in order to assess the viability of further investigations based on shell layer minor elements, isotopic or biochemical compositions. The results show that the shells seem to be well conserved, with microstructural patterns preserved down to sub-micrometric scales, and that some organic components are still present in situ. But faint taphonomic degradations affecting both mineral and organic components are nonetheless evidenced, such as the disappearance of organic envelopes surrounding crossed-foliated lamellae, combined with a partial recrystallization of the lamellae. Our results provide a solid case-study of the early stages of the diagenetic evolution of crossed-foliated shell layers. Moreover, they highlight the fact that extreme caution must be taken before using fossil shells for palaeoenvironmental or geochronological reconstructions. Without thorough investigation, the alteration patterns illustrated here would easily have gone unnoticed. However, these degradations are liable to bias any proxy based on the elemental, isotopic or biochemical composition of the shells. This study also provides significant data concerning human subsistence behavior: the presence of notches and the good preservation state of limpet shells (no dissolution/recrystallization, no bioerosion and no abrasion/fragmentation aspects) would attest that limpets were gathered alive with tools by Middle Palaeolithic (Aterian) populations in North Africa for consumption

Molecular Characterization of the Gastrula in the Turtle Emys orbicularis: An Evolutionary Perspective on Gastrulation

Due to the presence of a blastopore as in amphibians, the turtle has been suggested to exemplify a transition form from an amphibian- to an avian-type gastrulation pattern. In order to test this hypothesis and gain insight into the emergence of the unique characteristics of amniotes during gastrulation, we have performed the first molecular characterization of the gastrula in a reptile, the turtle Emys orbicularis. The study of Brachyury, Lim1, Otx2 and Otx5 expression patterns points to a highly conserved dynamic of expression with amniote model organisms and makes it possible to identify the site of mesoderm internalization, which is a long-standing issue in reptiles. Analysis of Brachyury expression also highlights the presence of two distinct phases, less easily recognizable in model organisms and respectively characterized by an early ring-shaped and a later bilateral symmetrical territory. Systematic comparisons with tetrapod model organisms lead to new insights into the relationships of the blastopore/blastoporal plate system shared by all reptiles, with the blastopore of amphibians and the primitive streak of birds and mammals. The biphasic Brachyury expression pattern is also consistent with recent models of emergence of bilateral symmetry, which raises the question of its evolutionary significance