Crystallographic characterization of the crossed lamellar structure in the bivalve Meretrix lamarckii using electron beam techniques

To understand the formation mechanism of crossed lamellar structures in molluskan shells, the crystallographic structural features in the shell of a bivalve, Meretrix lamarckii, were investigated using scanning electron microscopy, electron backscattered diffraction, and transmission electron microscopy with a focused ion beam sample preparation technique. Approximately 0.5 μm-thick lamellae (the second-order units) are piled up obliquely toward the growth direction to form the first-order unit and the obliquity is inverted between adjacent units along the shell thickness direction. The first-order units originate around the center of the shell, initially growing parallel to the shell and subsequently curving toward the inner or outer surfaces. The lamellae consist of aragonite granular and columnar layers, which group together to adopt the same crystal orientation forming crystallographic units (crystallites). Multiple {1 1 0} twins are common both in the granular and columnar layers. The crystallite c-axis is parallel to the columns and is inclined at angles 0–50° from the lamellar normal (dispersing among individual lamellae), toward the shell growth direction. Probably, the directions of the a- and b-axes are random in the lamellae, showing no specific orientation.     

Bending and branching of calcite laths in the foliated microstructure of pectinoidean bivalves occurs at coherent crystal lattice orientation

Foliated calcite is widely employed by some important pteriomorph bivalve groups as a construction material. It is made from calcite laths, which are inclined at a low angle to the internal shell surface, although their arrangement is different among the different groups. They are strictly ordered into folia in the anomiids, fully independent in scallops, and display an intermediate arrangement in oysters. Pectinids have particularly narrow laths characterized by their ability to change their growth direction by bending or winding, as well as to bifurcate and polyfurcate. Electron backscatter analysis indicates that the c-axes of laths are at a high, though variable, angle to the growth direction, and that the laths grow preferentially along the projection of an intermediate axis between two a-axes, although they can grow in any intermediate direction. Their main surfaces are not particular crystallographic faces. Analyses done directly on the lath surfaces demonstrate that, during the bending/branching events, all crystallographic axes remain invariant. The growth flexibility of pectinid laths makes them an excellent space-filling material, well suited to level off small irregularities of the shell growth surface. We hypothesize that the exceptional ability of laths to change their direction may be promoted by the mode of growth of biogenic calcite, from a precursor liquid phase induced by organic molecules.     

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.     

Geometrical and crystallographic constraints determine the self-organization of shell microstructures in Unionidae (Bivalvia: Mollusca)

Unionid shells are characterized by an outer aragonitic prismatic layer and an inner nacreous layer. The prisms of the outer shell layer are composed of single-crystal fibres radiating from spheruliths. During prism development, fibres progressively recline to the growth front. There is competition between prisms, leading to the selection of bigger, evenly sized prisms. A new model explains this competition process between prisms, using fibres as elementary units of competition. Scanning electron microscopy and X-ray texture analysis show that, during prism growth, fibres become progressively orientated with their three crystallographic axes aligned, which results from geometric constraints and space limitations. Interestingly transition to the nacreous layer does not occur until a high degree of orientation of fibres is attained. There is no selection of crystal orientation in the nacreous layer and, as a result, the preferential orientation of crystals deteriorates. Deterioration of crystal orientation is most probably due to accumulation of errors as the epitaxial growth is suppressed by thick or continuous organic coats on some nacre crystals. In conclusion, the microstructural arrangement of the unionid shell is, to a large extent, self-organized with the main constraints being crystallographic and geometrical laws.     

REMOTE BIOMINERALIZATION IN DIVARICATE RIBS OF STRIGILLA AND SOLECURTUS (TELLINOIDEA: BIVALVIA)

Deposits composed of aragonite prisms, which were formed afterthe outer shell layer, have been found at the posterior steepslopes of divaricate ribs in two species of Strigilla and anothertwo of Solecurtus. These prisms have their axes oriented perpendicularto the outer shell surface and differ in morphology from fibresof the surface-parallel composite prisms forming the outer shell.They display crystalline features indicating that, unlike crystalsforming the outer shell surface, their growth front was free,unconstrained by the mantle or periostracum. These particulardeposits are called free-growing prisms (FGPs). In these generathe periostracum is clearly not the substrate for biomineralizationand, upon formation, does not adhere to the steep slope of ribs,but detaches at the rib peak and reattaches towards the posterior,just beyond the foot of the posterior scarps of ribs. In thisway, a sinus or open space developed between the internal surfaceof the periostracum and the outer shell surface along each steeprib slope. These spaces could remain filled with extrapallialfluid after the mantle advances beyond that point during shellsecretion. FGPs grow within this microenvironment, out of contactwith the mantle. Other species with divaricate ribs do not developFGPs simply because the periostracum adheres tightly to both ribslopes (which are never so steep as in Solecurtus and Strigilla).FGPs constitute one of the rare cases of remote biomineralizationin which aragonite is produced and direct contact with the mantlenever takes place. (Received 22 November 1999; accepted 20 February 2000)     

Great spotted cuckoo eggshell microstructure characteristics can make eggs stronger

Obligate avian brood parasites lay stronger eggs than their hosts or non‐parasitic relatives because they are rounder and have a thicker eggshell. Additionally, some other characteristics of the brood parasitic eggshells related to their microstructure such as size and orientation of calcite crystal units could also contribute to generating even stronger shells. An eggshell microstructure formed by small randomly oriented calcite crystal units can increase the robustness of the eggshells of birds. Here, the eggshell microstructure of avian brood parasites as well as their hosts have been characterized in detail, using X‐ray diffraction analyses to estimate the size and degree of orientation of calcite crystal units making the eggshell. Specifically, the brood parasitic great spotted cuckoo Clamator glandarius and two hosts (jackdaws Corvus monedula and magpie Pica pica) and one non‐host species (the pigeon, Columba livia domestica) were considered. Calcite crystal of the eggshell of the brood parasitic species was smaller and more randomly oriented than those of the eggshells of non‐parasitic species, which suggest that eggshell microstructure would contribute to explain why parasitic eggs are more resistant to breakage than those of their hosts.     

On the nature of the opaque and translucent enamel regions of some macropodinae (Macropus giganteus,Wallabia bicolor and Peradorcas concinna)

Summary Teeth of three macropod species, M. giganteus, W. bicolor and P. concinna, have been studied using the techniques of light microscopy, scanning- and transmission-electron microscopy and hardness measurement. Light microscope observations showed that the teeth of these species had a translucent enamel region close to the dentine and an outer opaque enamel region at the tooth's surface. These regions were not related to the presence or absence of tubules which are a characteristic feature of marsupial enamel. Hardness tests showed that the opaque enamel was softer than the translucent enamel. Scanning electron microscope observations revealed that there was no correlation between any particular prism packing or orientation and the opaque and translucent enamel regions. Transmission electron microscope observations showed that the translucent enamel region consisted of well defined prisms and well packed, lath-like crystals, whereas the opaque enamel was disrupted by voids (which ranged in size from enlarged micropores to about 2 m in diameter in extreme cases) between crystals and some randomly oriented, loosely packed crystals. This disruption within the opaque enamel region was more common at prism boundaries but pockets of disrupted enamel were also found within prisms and interprismatic regions. The opacity of the enamel was caused by scattering of light from the voids. The ultrastructure of the opaque enamel region indicated that this region was hypomineralized; hardness tests and polarized light microscope observations were consistent with these results.     

Parataxonomic review of the Upper Cretaceous dinosaur eggshells belonging to the oofamily Megaloolithidae from India and Argentina

The eggshell oospecies from India and Argentina are compared and reviewed in detail. These eggshells resemble each other in having a nodular outer surface ornamentation and clearly arched growth lines of the shell units. Microstructurally, the eggshell oospecies belonging to the oofamily Megaloolithidae shows fan-like shell units, which are sharply separated from each other throughout the thickness of the eggshell and can be traced up to the surface of the eggshell. Comparisons between four oospecies from India and Argentina reveal three groupings, which show similarities between megaloolithids of both countries: (1) Megaloolithus jabalpurensis, M. matleyi and M. patagonicus; (2) M. cylindricus, M. rahioliensis and Tipo 1d; and (3) M.megadermus and Tipo 1e. The other two types of eggshell oospecies from India and Argentina show partially fused external nodes and shell units. As a result, growth lines enter into the adjacent shell units with a marked concavity. A new oogenus Fusioolithus have been erected due to fusion between shell units and tubospherulitic morphotype, which include two new oospecies F. baghensis and F. berthei. Till date, morphostructurally, a total of 15 eggshell oospecies belonging to different oofamilies have been recorded from India and seven oospecies from Argentina.     

THE STRUCTURE AND ORGANIZATION OF, AND THE RELATIONSHIP BETWEEN THE ORGANIC MATRIX AND THE INORGANIC CRYSTALS OF EMBRYONIC BOVINE ENAMEL

Electron microscope and electron diffraction studies of developing embryonic bovine enamel have revealed the organization of the organic matrix and the inorganic crystals. The most recently deposited inorganic crystals located at the ameloblast-enamel junction are thin plates, approximately 1300 A long, 400 A wide, and 19 A thick. During maturation of the enamel, crystal growth occurs primarily by an increase in crystal thickness. Statistical analyses failed to show a significant change in either the width or the length of the crystals during the period of maturation studied. Even in the earliest stages of calcification, the crystals are organized within the prisms so that their long axes (c-axes) are oriented parallel to the long axes of the prisms but randomly distributed about their long axes. With maturation of the enamel, the crystals become more densely packed and more highly oriented within the prisms. The organic matrix in decalcified sections of enamel is strikingly similar in its over-all organization to that of the fully mineralized tissue. When viewed in longitudinal prism profiles, the intraprismatic organic matrix is composed of relatively thin dense lines, approximately 48 A wide, which are relatively parallel to each other and have their fiber axes parallel to the long axes of the prisms within which they are located. Many of these dense lines, which have the appearance of thin filaments, are organized into doublets, the individual 48 A wide filaments of the doublets being separated by approximately 120 A. When observed in oblique prism profiles, the intraprismatic organic matrix is likewise remarkably similar in general orientation and organization to that of the fully mineralized tissue. Moreover, the spaces between adjacent doublets or between single filaments have the appearance of compartments. These compartments, more clearly visualized in cross- or near cross-sectional prism profiles, are oval or near oval in shape. Therefore, the appearance of the intraprismatic organic matrix (in longitudinal, oblique, and cross-sectional prism profiles) indicates that it is organized into tubular sheaths which are oriented with their long axes parallel to the long axes of the prisms in which they are located, but randomly oriented about their own long axes, an orientation again remarkably "blue printing" that of the inorganic crystals. The predominant feature of the walls of the tubular sheaths, when viewed in cross- or near cross-section, is that of continuous sheets, although in many cases closely packed dot-like structures of approximately 48 A were also observed, suggesting that the wall of the sheaths consists of a series of closely packed filaments. The 48 A wide dense lines (filaments) representing the width of the sheath wall were resolved into two dense strands when viewed in longitudinal prism profiles. Each strand was 12 A wide and was separated by a less electron-dense space 17 A wide. The intraprismatic organic matrix is surrounded by a prism sheath which corresponds in mineralized sections to the electron-lucent uncalcified regions separating adjacent prisms. Structurally, the prism sheaths appear to consist of filaments arranged in basket-weave fashion.     

Ab initio and molecular dynamics studies of solid β-HMX: effects of hydrostatic pressure and high temperature

Using first-principles density functional theory and classical molecular dynamics (MD), the structural, electronic and mechanical properties of the energetic material β-HMX have been studied. The crystal structure optimised by the local density approximation calculations compares reasonably with the experimental data. Electronic band structure and density of states indicate that β-HMX is an insulator with a band gap of 3.059 eV. The pressure effect on the crystal structure and physical properties has been investigated in the range of 0–40 GPa. The crystal structure and electronic properties change slightly as the pressure increases from 0 to 2.5 GPa; when the pressure is above 2.5 GPa, further increment of the pressure results in significant changes in crystal structure. There is a larger compression along the b-axis than along the a- and c-axes. Isothermal–isobaric MD simulations on β-HMX were performed in the temperature range of 5–400 K. Phase transition at 360 K, corresponding to a volume interrupt, was found. The computed thermal expansion coefficients show anisotropic behaviour with a slightly larger expansion along the b- and c-axes than along the a-axis. In the temperature range of 5–360 K, β-HMX possesses good plasticity and its stiffness decreases with increasing the temperature.     

Control of biofouling on pearl oysters Pinctada imbricata using wax and Chinese herbs

Abstract

Biofouling poses severe challenges to pearl oyster Pinctada imbricata culture in China, and controlling it is both labor- and capital-intensive. The antifouling properties of wax, and wax mixed with Chinese herbs, sprayed onto pearl oyster shell surfaces during peak biofouling seasons were evaluated. Pearl oysters coated with three wax treatments (plain wax, Chinaberry seed extract, Chinese honeylocust fruit extract) and a control (no treatment), were cultured in nets for up to 60?days. Mortality rate, fouling organism and pearl-oyster weights, and shell height are reported for individual oysters on each of six sampling dates. With the exception of oysters submerged for 12?days, all oysters were significantly affected by treatment type and submersion duration. Fouling weight increased more rapidly over time in the control-treatment oysters. Wax-based coatings deterred fouling-organism settlement on oysters for at least 2?months during the intensive fouling season, reducing mortality and not adversely effecting growth.     

Microstructure and growth of the lenses of schizochroal trilobite eyes

Lenses within the schizochroal eyes of phacopine trilobites are made principally of calcite, and characterization of them using light microscopy and high‐resolution electron imaging and diffraction has revealed an array of microstructural arrangements that suggest a common original pattern across the suborder. The low convexity lenses of Odontochile hausmanni and Dalmanites sp. contain calcite fibres termed trabeculae. The c axis of trabecular calcite lies parallel to the lens axis, and adjacent trabeculae are distinguished by small differences in their a axis orientations. Despite the common alignment, the boundaries between trabeculae cross‐cut the c axis as they fan out towards the lens base. Trabeculae are absent from the lens immediately beneath the visual surface, and instead, a radial fringe is present and is composed of micrometre‐thick sheets of calcite whose c axes are oriented at a low angle to the visual surface. High convexity lenses are more common than those of lower convexity among the species studied, and they have a much thicker radial fringe. Beneath this fringe, all of the lens calcite is oriented with its c axis parallel to the lens axis and it lacks trabeculae. We propose that both the high and low convexity lenses formed by rapid growth of calcite from a surface that migrated inwards from the cornea, and they may have had an amorphous calcium carbonate precursor. The trabeculae and radial fringes are unlikely to have had any beneficial effect on the transmission or focusing of light, but rather are the outcomes of an elegant solution to the problem of how to construct a biconvex lens from a crystalline solid.     

Selectively bred oysters can alter their biomineralization pathways,promoting resilience to environmental acidification

Commercial shellfish aquaculture is vulnerable to the impacts of ocean acidification driven by increasing carbon dioxide (CO₂) absorption by the ocean as well as to coastal acidification driven by land run off and rising sea level. These drivers of environmental acidification have deleterious effects on biomineralization. We investigated shell biomineralization of selectively bred and wild‐type families of the Sydney rock oyster Saccostrea glomerata in a study of oysters being farmed in estuaries at aquaculture leases differing in environmental acidification. The contrasting estuarine pH regimes enabled us to determine the mechanisms of shell growth and the vulnerability of this species to contemporary environmental acidification. Determination of the source of carbon, the mechanism of carbon uptake and use of carbon in biomineral formation are key to understanding the vulnerability of shellfish aquaculture to contemporary and future environmental acidification. We, therefore, characterized the crystallography and carbon uptake in the shells of S. glomerata, resident in habitats subjected to coastal acidification, using high‐resolution electron backscatter diffraction and carbon isotope analyses (as δ¹³C). We show that oyster families selectively bred for fast growth and families selected for disease resistance can alter their mechanisms of calcite crystal biomineralization, promoting resilience to acidification. The responses of S. glomerata to acidification in their estuarine habitat provide key insights into mechanisms of mollusc shell growth under future climate change conditions. Importantly, we show that selective breeding in oysters is likely to be an important global mitigation strategy for sustainable shellfish aquaculture to withstand future climate‐driven change to habitat acidification.     

Continuous and discontinuous growth in the otolith of Macruronus novaezelandiae (Merlucciidae: Teleostei)

Viewed by transmitted light, the lapillus and astericus otoliths Macruronus novaezelandiae (Merlucciidae) contain radial bands of similar width to the distances between steps on the surface of the otolith. The sagitta otolith has a multi-prismatic structure and shows differences in ultrastructure between its dorsal and ventral growth axes, as well as its sulcal (inward-facing) and anti-sulcal (outward-facing) parts. The ultrastructure of the sagitta shows that not all of the optical and etched checks in the central growth axis represent true discontinuities in the crystal growth of the otolith; they are the result of refraction around narrow optically active refractive bands. Microincrement growth along the dorsal prisms of the otolith from the primordium to the growing edge shows discontinuities in crystal growth at the boundary of the primordium and at the intersection of prisms. Parts of the ventral edge of the otolith show discontinuous crystal growth apparently caused by the physical growth restriction at the point at which the otolith is supported by the skull. Both the anti-sulcal and sulcal parts of the otolith often show discontinuities in the crystal structure alongside zones of continuous microincrement deposition, or evidence of continuous crystal growth, documenting simultaneous continuous and discontinuous growth in closely juxtaposed parts of the same otolith. © 1993 Wiley-Liss, Inc.     

Direct effects of physical stress can be counteracted by indirect benefits: oyster growth on a tidal elevation gradient

The paradigmatic gradient for intertidal marine organisms of increasing physical stress from low to high elevation has long served as the basis for using direct effects of duration of water coverage to predict many biological patterns. Accordingly, changes in potential feeding time may predict the direction and magnitude of differences between elevations in individual growth rates of sessile marine invertebrates. Oysters (triploid Crassostrea ariakensis) experimentally introduced at intertidal (MLW+0.05 m) and subtidal (MLW–0.25 m) elevations in racks provided a test of the ability to use duration of water coverage to predict changes in growth. During early-to-mid winter, a depression of 38–47% in shell growth of intertidal oysters matched the 36% reduction in available feeding time relative to subtidal oysters. In late winter as solar heating of exposed oysters increased, growth differences of 52–55% departed only slightly from the predicted 39%. In spring, however, duration of water coverage failed to predict even the correct direction of growth change with elevation as intertidal oysters grew 34% faster despite 39% less feeding time. Intense seasonal development of shell fouling by other suspension feeders like ascidians, mussels, and barnacles on subtidal (94% incidence) but not on aerially exposed intertidal (21–38% incidence) oysters may explain why duration of water cover failed to predict spring growth differences. Less intense fouling develops on intertidal oysters due to the physiological stress of aerial exposure on settlers, especially during higher temperatures and longer solar exposures of spring. Fouling by suspension feeders is known to reduce growth of the host through localized competition for food and added energetic costs. Thus, in springtime, indirect effects of aerial exposure providing a partial refuge from biological enemies overwhelmed direct effects of reduced duration of water coverage to reverse the expected pattern of slower intertidal growth of a marine invertebrate.     

Microtexture of larval shell of oyster,Crassostrea nippona: A FIB-TEM study

The initial formation and subsequent development of larval shells in marine bivalve, Crassostrea nippona were investigated using the FIB-TEM technique. Fourteen hours after fertilization (the trochophore stage), larvae form an incipient shell of 100–150 nm thick with a columnar contrast. Selected-area electron diffraction analysis showed a single-crystal aragonite pattern with the c-axis perpendicular to the shell surface. Plan-view TEM analysis suggested that the shell contains high density of {110} twins, which are the origin of the columnar contrast in the cross-sectional images. 72 h after fertilization (the veliger stage), the shell grows up to 1.2–1.4 μm thick accompanying an additional granular layer between the preexisting layer and embryo to form a distinctive two-layer structure. The granular layer is also composed of aragonite crystals sharing their c-axes perpendicular to the shell surface, but the crystals are arranged with a flexible rotation around the c-axes and not restricted solely to the {110} twin relation. No evidence to suggest the existence of amorphous calcium carbonate (ACC) was found through the observation. The well-regulated crystallographic properties found in the present sample imply initial shell formation probably via a direct deposition of crystalline aragonite.     

Model of the orientation symmetry of water molecules in biopolymers (by NMR spectra)

Spectra of proton and deuteron magnetic resonance of water molecules in oriented biopolymer samples (collagen, DNA, cellulose) are interpreted on the basis of a model of molecular Schottki-type diffusion. The presence of narrow splittings are shown to be the result of slight distortions of “ideal” symmetry of orientations of p-p and O-D vectors, corresponding to the point symmetry group of a tetrahedron, one of whose C₃-axes is oriented along the fiber axis.     

Structure of shells from eggs of kinosternid turtles

Shells from eggs of five species of kinosternid turtle (Sternotherus minor, Kinosternon flavescens, K. baurii, K. Hirtipes, and K. alamosae) were examined with light and scanning electron microscopy. Except for possible differences among species in thickness of eggshells, structure of shells from all eggs was similiar. In general, kinosternid turtles lay eggs having a rigid calcareous layer composed of calcium carbonate in the form of aragonite. The calcareous layer is organized into individual shell units with needlelike crystallites radiating from a common center. Most of the thickness of the eggshell is attributable to the calcareous layer, with the fibrous shell membrane comprising only a small fraction of shell thickness. Pores are found in the calcareous layer, but they are not numereous. The outer surface of the eggshells is sculptured and may have a thick, organic layer in places. The outer surface of the shell membrane of decalcified eggshells is studded with spherical cores which presumably nucleate growth of shell units during shell formation. The shell membrane detaches from eggs incubated to hatching, carrying with it remnants of the calcareous layer. Such changes in shell structure presumably reflect withdrawal of calcium from the eggshell by developing embryos.     

Crystallization and preliminary X-ray crystallographic analysis of ImmE7 protein of colicin E7

The ImmE7 protein, which can bind specifically to the DNase colicin E7 and neutralize its bactericidal activity, has been purified and crystallized in two different crystal forms by vapor diffusion method. The orthorhombic crystals belong to space group I222 or I2₁2₁2₁ and have unit cell dimensions a = 75.1 Å, b = 50.5 Å, and c = 45.4 Å. The second form is monoclinic space group P2₁ with ceil dimensions a = 29.3 Å, b = 102.7 Å, c = 53.0 Å and β = 91.5°. The orthorhombic crystals diffract to 1.8 Å resolution, and are suitable for high-resolution X-ray analysis. © 1995 Wiley-Liss, Inc.     

Organization and mode of secretion of the granular prismatic microstructure of Entodesma navicula (Bivalvia: Mollusca)

The term homogeneous has been applied to molluscan microstructures that lack a readily discernible microstructure and as a result, it has become rather a ‘dustbin’ term, covering a multitude of unrelated finely crystalline textures. Here we investigate in detail the outer ‘homogeneous’ layer of the lyonsiid bivalve Entodesma navicula. The apparently equigranular crystals (up to 10 µm) are, in fact, short prisms which grow in a dense organic matrix with their c‐axes and fibre axes coincident, perpendicular to the growth surface. These prisms are distinct from the aragonitic prisms grown by other bivalves in both their morphology and their mode of growth and so we propose the term granular prismatic microstructure. The organic content of granular prisms (7.4%) is the highest yet recorded for any molluscan microstructure and it is apparent that the short prisms have grown within a gel‐filled space. Although this high organic content is likely to make the microstructure metabolically expensive to produce, it has the benefit of making the valves very flexible. This may be advantageous in the intertidal zone inhabited by E. navicula by allowing a tight seal between the valves.