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Bivalves have evolved a range of complex shell forming mechanisms that are reflected by their incredible diversity in shell mineralogy and microstructures. A suite of proteins exported to the shell matrix space plays a significant role in controlling these features, in addition to underpinning some of the physical properties of the shell itself. Although, there is a general consensus that a minimum basic protein tool kit is required for shell construction, to date, this remains undefined. In this study, the shell matrix proteins (SMPs) of four highly divergent bivalves (The Pacific oyster, Crassostrea gigas; the blue mussel, Mytilus edulis; the clam, Mya truncata, and the king scallop, Pecten maximus) were analyzed in an identical fashion using proteomics pipeline. This enabled us to identify the critical elements of a “basic tool kit” for calcification processes, which were conserved across the taxa irrespective of the shell morphology and arrangement of the crystal surfaces. In addition, protein domains controlling the crystal layers specific to aragonite and calcite were also identified. Intriguingly, a significant number of the identified SMPs contained domains related to immune functions. These were often are unique to each species implying their involvement not only in immunity, but also environmental adaptation. This suggests that the SMPs are selectively exported in a complex mix to endow the shell with both mechanical protection and biochemical defense.

Molluscs fabricate shells of incredible diversity and complexity by localized secretions from the dorsal epithelium of the mantle. Although distantly related molluscs express remarkably different secreted gene products, it remains unclear if the evolution of shell structure and pattern is underpinned by the differential co-option of conserved genes or the integration of lineage-specific genes into the mantle regulatory program. To address this, we compare the mantle transcriptomes of 11 bivalves and gastropods of varying relatedness. We find that each species, including four Pinctada (pearl oyster) species that diverged within the last 20 Ma, expresses a unique mantle secretome. Lineage- or species-specific genes comprise a large proportion of each species’ mantle secretome. A majority of these secreted proteins have unique domain architectures that include repetitive, low complexity domains (RLCDs), which evolve rapidly, and have a proclivity to expand, contract and rearrange in the genome. There are also a large number of secretome genes expressed in the mantle that arose before the origin of gastropods and bivalves. Each species expresses a unique set of these more ancient genes consistent with their independent co-option into these mantle gene regulatory networks. From this analysis, we infer lineage-specific secretomes underlie shell diversity, and include both rapidly evolving RLCD-containing proteins, and the continual recruitment and loss of both ancient and recently evolved genes into the periphery of the regulatory network controlling gene expression in the mantle epithelium.

Reductions to seawater pH challenge the shell integrity of marine calcifiers. Many molluscs have an external organic layer (the periostracum) that limits exposure of underlying shell to the outside environment, which could potentially help combat shell dissolution under corrosive seawater conditions. We tested this hypothesis in adult California mussels, Mytilus californianus . We quantified shell dissolution rates as a function of periostracum cover across three levels of reduced pH (7.7, 7.5, and 7.4 on the total scale). Since periostracum can also be eroded over time, we additionally conducted a first-pass examination of whether differing surface textures induced by abrasional processes might influence dissolution rates. We contextualized this set of experiments with measurements of mussel periostracum cover in multiple intertidal habitats. Our results indicate a threefold reduction in shell dissolution rate as periostracum cover increases from 10 to 85% of shell surface area. Dissolution was higher in lower-pH treatments and in treatments where periostracum removal resulted in shells with rougher surface texture, potentially due to increased microtopographic surface area of underlying shell exposed to corrosive seawater. Periostracum loss in the field was greater for mussels at higher shoreline elevations and in sunnier locations, where heat, ultraviolet radiation, and desiccation at low tide may weaken attachment of the periostracum to the shell and. These findings highlight the potential for protective structures of marine organisms to help confront increasingly acute global environmental stressors.

Helicoidal formations often appear in natural microstructures such as bones and arthropods exoskeletons. Named Bouligands after their discoverer, these structures are angle-ply laminates that assemble from laminae of chitin or collagen fibers embedded in a proteinaceous matrix. High resolution electron microscope images of cross-sections through scorpion claws are presented here, uncovering structural features that are different than so-far assumed. These include in-plane twisting of laminae around their corners rather than through their centers, and a second orthogonal rotation angle which gradually tilts the laminae out-of-plane. The resulting Bouligand laminate unit (BLU) is highly warped, such that neighboring BLUs are intricately intertwined, tightly nested and mechanically interlocked. Using classical laminate analysis extended to laminae tilting, it is shown that tilting significantly enhances the laminate flexural stiffness and strength, and may improve toughness by diverting crack propagation. These observations may be extended to diverse biological species and potentially applied to synthetic structures. Helicoids are common structures found in many structural biological materials. Here, the authors report on a study of helicoids in the claws of scorpions and report different microstructures to what have previously been reported which have implications in materials stiffness, strength and toughness.

Climate change is a global environmental threat, directly affecting biodiversity. Terrestrial gastropods are particularly susceptible to alterations in temperature and humidity and have develop morph-physiological and behavioural adaptations in this regard. Shell colour polymorphism and its potential implication for thermoresistance constitute an unexplored field in Neotropical land snails. The variation in shell colour luminance is characterized in the threatened endemic Eastern Cuban tree snails Polymita picta and Polymita muscarum using digital tools; being able to discriminate shell luminance between colour morphs for both species, under different image-taking conditions. For P . muscarum , the albino morph presented the highest luminance values (152.7 ± 0.4); while the lowest values correspond to the brown morph with dark bands (112.9 ± 0.8). Otherwise, for P . picta , the morphs showing the highest luminance were yellow with a pink sutural band (112.8 ± 7.1) and pale yellow (112.6 ± 10.3) and the lowest luminance corresponded to the black morph (44.5 ± 1.2). The presence of dark bands decreased the luminance values regardless of their position in the shell, the morph and the species analysed. In general, the shells of P . muscarum have higher luminance than those of P . picta . Luminance variations demonstrate the ’indiscrete’ nature of this trait and highlight the complex interactions between evolutionary mechanisms and shell color polymorphism in Polymita . This supports the hypothesis that colour has adaptive value for thermoregulation, encompassing not only the background colour but also the coloration of the bands. The differences in the shell luminance in both species suggest a correlation with the geographical distribution and corresponding habitats. Based on our findings, yellowish morphs will be more resistant to future climatic conditions in their respective habitats on the island.

The dissolution of the delicate shells of sea butterflies, or pteropods, has epitomised discussions regarding ecosystem vulnerability to ocean acidification over the last decade. However, a recent demonstration that the organic coating of the shell, the periostracum, is effective in inhibiting dissolution suggests that pteropod shells may not be as susceptible to ocean acidification as previously thought. Here we use micro-CT technology to show how, despite losing the entire thickness of the original shell in localised areas, specimens of polar species Limacina helicina maintain shell integrity by thickening the inner shell wall. One specimen collected within Fram Strait with a history of mechanical and dissolution damage generated four times the thickness of the original shell in repair material. The ability of pteropods to repair and maintain their shells, despite progressive loss, demonstrates a further resilience of these organisms to ocean acidification but at a likely metabolic cost. Sea butterflies, or pteropods, are often presented as being at threat from ocean acidification on account of their fragile shells being susceptible to dissolution. Here the authors show that pteropods are able to perform extensive repair to damaged shells, suggesting they may not be as vulnerable as previously thought.

Molluscan shells are a classic model system to study formation–structure–function relationships in biological materials and the process of biomineralized tissue morphogenesis. Typically, each shell consists of a number of highly mineralized ultrastructures, each characterized by a specific 3D mineral–organic architecture. Surprisingly, in some cases, despite the lack of a mutual biochemical toolkit for biomineralization or evidence of homology, shells from different independently evolved species contain similar ultrastructural motifs. In the present study, using a recently developed physical framework, which is based on an analogy to the process of directional solidification and simulated by phase-field modeling, we compare the process of ultrastructural morphogenesis of shells from 3 major molluscan classes: A bivalve Unio pictorum, a cephalopod Nautilus pompilius, and a gastropod Haliotis asinina. We demonstrate that the fabrication of these tissues is guided by the organisms by regulating the chemical and physical boundary conditions that control the growth kinetics of the mineral phase. This biomineralization concept is postulated to act as an architectural constraint on the evolution of molluscan shells by defining a morphospace of possible shell ultrastructures that is bounded by the thermodynamics and kinetics of crystal growth.

The function-optimized properties of biominerals arise from the hierarchical organization of primary building blocks. Alteration of properties in response to environmental stresses generally involves time-intensive processes of resorption and reprecipitation of mineral in the underlying organic scaffold. Here, we report that the load-bearing shells of the brachiopod Discinisca tenuis are an exception to this process. These shells can dynamically modulate their mechanical properties in response to a change in environment, switching from hard and stiff when dry to malleable when hydrated within minutes. Using ptychographic X-ray tomography, electron microscopy and spectroscopy, we describe their hierarchical structure and composition as a function of hydration to understand the structural motifs that generate this adaptability. Key is a complementary set of structural modifications, starting with the swelling of an organic matrix on the micron level via nanocrystal reorganization and ending in an intercalation process on the molecular level in response to hydration. Bioinspired materials require an understanding of how biomaterials achieve the materials properties. Here, the authors report on the load-bearing shell of Discinisca tenuis and explore how hydration changes the dry shell from hard and stiff to soft and flexible within minutes by reorganisation caused by organic matrix swelling.

Biological systems have a remarkable capability of synthesizing multifunctional materials that are adapted for specific physiological and ecological needs. When exploring structure–function relationships related to multifunctionality in nature, it can be a challenging task to address performance synergies, trade-offs, and the relative importance of different functions in biological materials, which, in turn, can hinder our ability to successfully develop their synthetic bioinspired counterparts. Here, we investigate such relationships between the mechanical and optical properties in a multifunctional biological material found in the highly protective yet conspicuously colored exoskeleton of the flower beetle, Torynorrhina flammea. Combining experimental, computational, and theoretical approaches, we demonstrate that a micropillar-reinforced photonic multilayer in the beetle’s exoskeleton simultaneously enhances mechanical robustness and optical appearance, giving rise to optical damage tolerance. Compared with plain multilayer structures, stiffer vertical micropillars increase stiffness and elastic recovery, restrain the formation of shear bands, and enhance delamination resistance. The micropillars also scatter the reflected light at larger polar angles, enhancing the first optical diffraction order, which makes the reflected color visible from a wider range of viewing angles. The synergistic effect of the improved angular reflectivity and damage localization capability contributes to the optical damage tolerance. Our systematic structural analysis of T. flammea’s different color polymorphs and parametric optical and mechanical modeling further suggest that the beetle’s microarchitecture is optimized toward maximizing the first-order optical diffraction rather than its mechanical stiffness. These findings shed light on material-level design strategies utilized in biological systems for achieving multifunctionality and could thus inform bioinspired material innovations.

Mollusks have developed a broad diversity of shelled structures to protect against challenges imposed by biological interactions(e.g., predation) and constraints (e.g., p C O 2 -induced ocean acidification and wave-forces). Although the study of shell biomechanical properties with nacreous microstructure has provided understanding about the role of shell integrity and functionality on mollusk performance and survival, there are no studies, to our knowledge, that delve into the variability of these properties during the mollusk ontogeny, between both shells of bivalves or across the shell length. In this study, using as a model the intertidal mussel Perumytilus purpuratus to obtain, for the first time, the mechanical properties of its shells with nacreous microstructure; we perform uniaxial compression tests oriented in three orthogonal axes corresponding to the orthotropic directions of the shell material behavior (thickness, longitudinal, and transversal). Thus, we evaluated whether the shell material’s stress and strain strength and elastic modulus showed differences in mechanical behavior in mussels of different sizes, between valves, and across the shell length. Our results showed that the biomechanical properties of the material building the P. purpuratus shells are symmetrical in both valves and homogeneous across the shell length. However, uniaxial compression tests performed across the shell thickness showed that biomechanical performance depends on the shell size (aging); and that mechanical properties such as the elastic modulus, maximum stress, and strain become degraded during ontogeny. SEM observations evidenced that compression induced a tortuous fracture with a delamination effect on the aragonite mineralogical structure of the shell. Findings suggest that P. purpuratus may become vulnerable to durophagous predators and wave forces in older stages, with implications in mussel beds ecology and biodiversity of intertidal habitats.