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Few studies have considered the capabilities of gastropods living in minerally-deficient acidified coastal waters to compensate for outer shell corrosion or compromised growing edge shell production. We compared inner shell thickening between pristine shells (control) and corroded shells (experiment) of two related intertidal neritid gastropod species from reduced salinity and acidified environments. We predicted that the rocky-shore, Nerita chamaeleon, which has greater access to shell building biomineralization substrates, should better control shell thickness than the estuarine, Neripteron violaceum. Accordingly, N. chameleon was found to compensate perfectly for variation in the thickness of the outer calcitic blocky layer (BL). Optimal shell thickness (OST) was maintained by selective reabsorption of the aperture ridge of the distal shell (aragonitic crossed-lamellar layer, CL) and by increased internal deposition of proximal (older) shell (aragonitic protocrossed lamellar, PCL). Despite greater exposure to acidification and hyposalinity, N. violaceum showed no significant compensatory shell thickening. These findings reveal that shell thickening capability may vary greatly among intertidal gastropods and that this may be constrained by environmental biomineralization substrate availability. Such environmentally-related responses carry implications for predicted future reductions in coastal water pH and salinity., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

Biological tissues may exhibit graded heterogeneities in structure and mechanical properties that are crucial to their function. One biological structure that shows variation in both structure and function is the molluscan radula: the organ comprises a chitinous membrane with embedded teeth and serves to process and gather food. The tooth morphologies had been well studied in the last decades, but the mechanical properties of the teeth are not known for the vast majority of molluscs. This knowledge gap restricts our understanding of how the radula is able to act effectively on a target surface whilst simultaneously resisting structural failure. Here we employed nanoindentation technique to measure mechanical properties (hardness and Young's modulus) on distinct localities of individual radular teeth from 24 species of African paludomid gastropods. These species have distinct ecological niches as they forage on algae on different feeding substrates. A gradual distribution of measured properties along the teeth was found in species foraging on solid or mixed feeding substrates, but soft substrate feeders exhibit teeth almost homogeneous in their biomechanical properties. The presence or absence of large-scale gradients in these taenioglossan teeth could directly be linked with their specific function and in general with the species ecology, whereas the radular tooth morphologies do not always and fully reflect ecology. STATEMENT OF SIGNIFICANCE: African Lake Tanganyika is well known for harbouring endemic and morphologically distinct genera. Its paludomid gastropods form a flock of high interest because of its diversity. As they show distinct radular tooth morphologies hypotheses about potential trophic specializations had always been at hand. Here we evaluated the mechanical properties Young's modulus and hardness of 9027 individual teeth from 24 species along the tooth by nanoindentation and related them with the gastropods' specific feeding substrate. We find that hard substrate feeders have teeth that are hard at the tips but much less stiff at the base and thus heterogeneous with respect to material properties, whereas soft substrate feeders have teeth that are flexible and homogenous with respect to material properties., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2021 Acta Materialia Inc. Published by Elsevier Ltd. All rights reserved.)

Recent technical innovations are revealing surprising patterns in mollusc shell pigmentation, such as an unexpectedly modest role for melanins and rapid divergences in the mix of pigments used to achieve similar colour patterns. The elucidation of the molecular genetic basis of shell pigmentation has been slow, probably because of the high genome complexity of gastropods and bivalves. Recent work within the old field of evolutionary ecology of shell pigmentation allows a greater role for the analysis of large-geographic-scale patterns (sometimes employing citizen-science data), as well as experimental field studies. However, the field remains dominated by land snails as model organisms, while colour pattern evolution in marine gastropods and bivalves, particularly those not exposed to visual predators, remains mysterious., (Copyright © 2020 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Ocean acidification is mainly being monitored using data loggers which currently offer limited coverage of marine ecosystems. Here, we trial the use of gastropod shells to monitor acidification on rocky shores. Animals living in areas with highly variable pH (8.6-5.9) were compared with those from sites with more stable pH (8.6-7.9). Differences in site pH were reflected in size, shape and erosion patterns in Nerita chamaeleon and Planaxis sulcatus. Shells from acidified sites were shorter, more globular and more eroded, with both of these species proving to be good biomonitors. After an assessment of baseline weathering, shell erosion can be used to indicate the level of exposure of organisms to corrosive water, providing a tool for biomonitoring acidification in heterogeneous intertidal systems. A shell erosion ranking system was found to clearly discriminate between acidified and reference sites. Being spatially-extensive, this approach can identify coastal areas of greater or lesser acidification. Cost-effective and simple shell erosion ranking is amenable to citizen science projects and could serve as an early-warning-signal for natural or anthropogenic acidification of coastal waters., (Copyright © 2019. Published by Elsevier B.V.)

Scanning electron microscopy, X-ray diffraction and Fourier transformed infrared spectroscopy have been used to characterize the microstructure and instrumented microindentation for the determination of the mechanical properties of Charonia Lampas Lampas shell. Both elastic modulus and hardness are found to be dependent on the texture of the three distinct layers. From the analysis of load-depth curves, the shell exhibits small viscoelastic behaviour at low indentation loads and mainly elastoplastic behaviour at higher loads. These phenomena were attributed to the influence of the organic matter present in the shell. Both elastic modulus and hardness are found to be load-dependent in each layer in relation to their microstructure and, accordingly, to the anisotropy of the predominant mineral part. At a macroscopic scale, this tendency is explained by using a rule of mixture and jointly by the anisotropy of the aragonite. The Bull and Page model is subsequently applied to the hardness variation in order to compute the macrohardness which is the characteristic hardness number of a material and the hardness parameter related to the indentation size effect. This model describes well the experimental results for the relative higher depths, and deviates for the small depths due to the effect of the viscoelastic behaviour which then requires a more appropriate model to describe this phenomenon., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Through controlled exposure to tributyltin (TBT), the effects of season, size, and population origin were evaluated on imposex incidence in Stramonita brasiliensis. Four experiments were carried out with organisms collected from three different sites on the Brazilian coast (Torres - Rio Grande do Sul, Farol de Santa Marta - Santa Catarina, and Aracruz - Espírito Santo). S. brasiliensis were anesthetized, sexed, measured, classified by size in small (< 30 mm), medium (≥ 30 to ≤ 40 mm), or big (> 40 mm) and injected in the foot muscle with 0.5 μg g -1 of TBT. Organisms were maintained during one month in aquariums with clear marine water under controlled laboratory conditions. One month after injection, the imposex assessments showed that the population from the southeast (tropical) region was more sensitive to TBT than the population from the south of Brazil (the temperate region). A greater sensitivity in the small and medium categories was observed. Females were also more susceptible to TBT when exposed during their reproductive period. Thus, the present study highlighted the need to ensure that intrinsic biological factors related to organisms are considered in such biomonitoring studies to avoid misinterpretation of results., (Copyright © 2018 Elsevier Inc. All rights reserved.)

Gastropods shells have evolved to resist the threat of increasingly stronger predators that smash, peal, and crush their shells. Their shells are most commonly constructed from a crossed lamellar microstructure, which consists of an exquisitely architected arrangement of aragonitic mineral and organic encompassing at least four orders of hierarchy. It is this careful control of mineral and organic placement within the entire crossed lamellar structure that yields a four-order of magnitude increase in fracture toughness versus abiotic aragonite. We investigated the effect of differing microstructural orientations on their influence of inter-3rd order lamellar fracture behavior using nanoindentation from the inner layer of the Strombus gigas shell. We observed a significant influence of lamella (plank) orientation and nanoindenter probe on the mechanical properties. The ±45° arrangement of mineral planks found within biological crossed lamellar composites provides a significant enhancement of isotropic resistance to penetration by sharp objects such as jaws and claws. In addition, the ±45° arrangement is able to resist higher loads before failure. This combination of features from the crossed lamellar architecture helped enable species with this shell structure to survive predation for hundreds of millions of years and will also help provide insights into designs of future generations of composites., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Metallic tungsten (W) was initially assumed to be environmentally benign and a green alternative to lead. However, subsequent investigations showed that fishing weights and munitions containing elemental W can fragment and oxidize into complex monomeric and polymeric tungstate (WO4) species in the environment; this led to increased solubility and mobility in soils and increased bioaccumulation potential in plant and animal tissues. Here we expand on the results of our previous research, which examined tungsten toxicity, bioaccumulation, and compartmentalization into organisms, and present in this research that the bioaccumulation of W was related to greater than 50% reduction in the mechanical properties of the snail (Otala lactea), based on depth-sensing nanoindentation. Synchrotron-based X-ray fluorescence maps and X-ray diffraction measurements confirm the integration of W in newly formed layers of the shell matrix with the observed changes in shell biomechanical properties, mineralogical composition, and crystal orientation. With further development, this technology could be employed as a biomonitoring tool for historic metals contamination since unlike the more heavily studied bioaccumulation into soft tissue, shell tissue does not actively eliminate contaminants., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

The morphology of gastropod shells has been a focus of analyses in ecology and evolution. It has recently emerged as an important issue in developmental biology, thanks to recent advancements in molecular biological techniques. The growing tube model is a theoretical morphological model for describing various coiling patterns of molluscan shells, and it is a useful theoretical tool to relate local tissue growth with global shell morphology. However, the growing tube model has rarely been adopted in empirical research owing to the difficulty in estimating the parameters of the model from morphological data. In this article, I solve this problem by developing methods of parameter estimation when (1) 3D Computed Tomography (CT) data are available and (2) only 2D image data (such as photographs) are available. When 3D CT data are available, the parameters can be estimated by fitting an analytical solution of the growing tube model to the data. When only 2D image data are available, we first fit Raup׳s model to the 2D image data and then convert the parameters of Raup׳s model to those of the growing tube model. To illustrate the use of these methods, I apply them to data generated by a computer simulation of the model. Both methods work well, except when shells grow without coiling. I also demonstrate the effectiveness of the methods by applying the model to actual 3D CT data and 2D image data of land snails. I conclude that the method proposed in this article can reconstruct the coiling pattern from observed data., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Mollusk shells contain a plethora of information on past climate variability. However, only a limited toolkit is currently available to reconstruct such data from the shells. The environmental data of some proxies (e.g. Sr/Ca ratios) is obscured by physiological effects, whereas other proxies, such as δ(18)O, simultaneously provide information on two or more different environmental variables. The present study investigates whether microstructures of the freshwater gastropod Viviparus viviparus provide an alternative means to reconstruct past water temperature. Cold and highly variable temperature regimes resulted in the precipitation of highly unordered first-order lamellae of simple crossed-lamellar (XLM) structures if new shell formed from scratch. However, during stable and warm conditions, well-ordered first-order lamellae were laid down irrespective of pre-existing shell material. Homogeneous first-order lamellae also formed during times of cold and highly variable temperatures if the new shell was deposited onto existing shell material with well-ordered first-order lamellae. The growth front seems to contain instructions for building specific microstructure variants, irrespective of environmental conditions. However, if this template is missing, the animal forms a deviating microstructure. Under extremely stressful situations (e.g. removal from habitat, calcein staining, extreme temperature shifts), the gastropod precipitates an evolutionarily older microstructure (irregular simple prisms) rather than XLM structures. These shell portions were macroscopically described as disturbance lines. In addition, repetitive, presumably periodic growth patterns were observed, which consisted of gradually changing third-order lamellae between consecutive faint, organic-rich growth lines. These growth patterns were probably controlled by intrinsic biological clocks and exhibited a two-daily periodicity. The results of this study may provide the basis for using changes in the microstructure of shell sections as a new sensor (environmental proxy) for past water temperature., (Copyright © 2014 Acta Materialia Inc. Published by Elsevier Ltd. All rights reserved.)

The contributions of mesolayers, organic interlamellar layers and nanoasperities/mineral bridges to the strength of nacre from red abalone (Haliotis rufescens) shell nacre are investigated. Samples were demineralized and deproteinized to separate the organic and mineral components, respectively. Tensile tests were performed on both the isolated organic constituent and the isolated mineral. The strength of the isolated organic component suggests that growth bands play an important role in the mechanical behavior as they are thick regions of protein that are a significant fraction (∼0.4) of the total organic content. The thickness variation of the nacre tablets was measured and found to be a small fraction of the mean tablet thickness (0.568μm); the standard deviation is 26nm, indicating that the wedge mechanism of toughening does not operate in the nacre investigated. Results obtained from the isolated mineral validate the importance of the organic constituent as the mechanical properties decline greatly when the organic component is removed. The results presented herein add to the understanding of the mechanical response of the organic interlayers and growth bands and their effect on the toughness of the abalone nacre., (Copyright © 2013 Acta Materialia Inc. Published by Elsevier Ltd. All rights reserved.)

We histologically examined normal differentiation and development of genital tract and gonad in the ivory shell Babylonia japonica (Buccinidae) to determine whether the formation of male-type genitalia in imposex-exhibiting females mimics the normal development of male genitalia in prosobranch gastropods. We used a wild-caught 2-year-old specimen and laboratory-reared juveniles aged 0-24 months. Gonad differentiation was unclear before age 14 months, but progressed after 16 months. Both sexes had complete genital tract and mature gonad at 20 months. However, differentiation and development occurred earlier in females than in males. Development of genital tract preceded gonad differentiation. Vas deferens morphogenesis in males resembled that in imposex-exhibiting females. These findings help to understand the morphogenesis of genital tract and gonad in prosobranch gastropods and will contribute to more in-depth studies of the mode of action of organotin compounds such as TBT and TPhT in imposex development in female prosobranch gastropods., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

The characteristics of the development of male genitalia (penis and vas deferens) in imposex-exhibiting female rock shells, Thais clavigera, were histologically examined using specimens from a wild population and tributyltin (TBT)-exposed females in the laboratory. A variety of vas deferens morphogenesis patterns were observed in wild female T. clavigera, and the characteristics were summarized. The immature vas deferens at an initial stage, however, was only observed beneath or behind the penis, and no vas deferens was observed close to the vaginal opening (i.e., vulva) of the capsule gland in TBT-exposed females, which was different from the characteristics of vas deferens formation observed in wild females. Taking into consideration both the observed results from wild female specimens and from TBT-exposed females in the laboratory, the vas deferens sequence (VDS) index for T. clavigera was proposed as VDS 1-6., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

Sea slugs (Gastropoda: Opisthobranchia) are characterized by extensive morphological homoplasy. In particular, reduced or absent shells are predominant throughout the group. This trend towards shell loss has resulted in a poor fossil record. DNA-based phylogenies have been helpful in improving our understanding of the evolution of this group and major clades are emerging. We report 13 new complete opisthobranch mitochondrial genomes that provide robust support for some of these emerging nodes. We name three new clades within the Opisthobranchia, the Actopleura (Acteonoidea plus Nudipleura), Placoesophaga (Cephalaspidea plus Anaspidea), and Siphoglossa (Sacoglossa plus the Siphonaria). Finally we use molecular clock dating that suggests an earlier opisthobranch divergence than previously reported. The implications of this evolutionary scenario are discussed., (Copyright © 2011 Elsevier B.V. All rights reserved.)

The shear strength of the interface between tiles of aragonite in the nacre of red abalone Haliotis rufescens was investigated through mechanical tensile and shear tests. Dog-bone shaped samples were used to determine the tensile strength of nacre when loaded parallel to the plane of growth; the mean strength was 65 MPa. Shear tests were conducted on a special fixture with a shear gap of 200 microm, approximately 100 microm narrower than the spacing between mesolayers. The shear strength is found to be 36.9+/-15.8 MPa with an average maximum shear strain of 0.3. Assuming the majority of failure occurs through tile pull-out and not through tile fracture, the tensile strength can be converted into a shear strength of 50.9 MPa. Three mechanisms of failure at the tile interfaces are discussed: fracture of mineral bridges, toughening due to friction created through nanoasperities, and toughening due to organic glue. An additional mechanism is fracture through individual tiles.

In this study we present the results of an intersex survey of Littorina littorea along the Belgian coast. Levels of female intersex and sterility were determined to assess TBT related adverse effects. In addition, we determined the levels of male penis shedding and trematode infestation and investigated the morphology of the shell. Significant differences were found for all these variables which clearly differentiated periwinkles from Zeebrugge (B2) from those at other locations. Intersex index (ISI) values were relatively low (i.e. 0.00-0.39), except at B2 where they ranged up to 3.52, the highest value ever reported in literature. Consequently, female reproductive impairment at B2 was severe. Indeed, up to 95% of female periwinkles were sterile at B2. In addition, 61% of the male periwinkles had shed their penis. Furthermore, no trematode infestation could be detected at B2 and specimens from this location had the largest and heaviest shells, which may be related to population demography and/or a different use of energy budgets.

Imposex in neogastropods was used to determine the relative TBT distribution in the Gulf of Thailand in 2006. To identify the imposex prevalence, 8757 specimens, belonging to 22 species from five families of neogastropods, were collected at 56 sites in 13 sample areas. These areas were located between the Bight of Bangkok in the north and the Malaysian border in the south. A contamination model was developed to compensate for differences in TBT sensitivity among species and to make comparisons among areas. At every area imposex was found in neogastropods. The highest incidence was in the east side of the Bight of Bangkok off Si Racha and Pattaya and in the southern part around Pattani. The same areas showed the highest frequency of imposex in 1996. While the frequency of imposex appeared to have slightly decreased in these areas, increases were found elsewhere. In Pattani Province this could be related to dumping of highly polluted harbour sediments in the shallow coastal area. The overall frequency of imposex in the Gulf of Thailand significantly increased from 1996 to 2006.

Although poorly reported in the scientific literature, acidic waters characterize many South East Asian estuaries. The observation of shell dissolution in a typically marine gastropod whelk (Thais) prompted investigation into determining relationships between shell properties of this whelk and the water chemistry (including pH) of the Sungai Brunei estuary (Borneo) in which it occurs. Shell weight, shell length and topographical shell features were determined for populations of Thais gradata distributed along a gradient of pH and salinity ranging between 5.78 and 8.3 pH units, and 3.58 and 31.2psu. Shell weight varied independently of the co-varying acidity, salinity and calcium levels experienced. In contrast, shell length and a semi-quantitative variable based on shell sculpturing (shell erosion rank, SER) were significantly correlated with these water chemistry variables. This study brings attention to the potential use of estuarine organisms and systems in investigating current marine acidification questions.

Strombus gigas (conch) shells and Tridacna gigas (Giant clam) shells have dense, tailored structures that impart excellent mechanical properties to these shells. In this investigation, conch and clam seashells were converted to hydroxyapatite (HAP) by a hydrothermal method at different temperatures and for different conversion durations. Dense HAP structures were created from these shells throughout the majority of the samples at the relative low temperature of approximately 200 degrees C. The average fracture stress was found to be approximately 137-218MPa for partially converted conch shell samples and approximately 70-150MPa for original and converted clamshell samples, which is close to the mechanical strength of compact human bone. This indicates that the converted shell samples can be used as implants in load-bearing cases. In vivo tests of converted shell samples were performed in rat femoral defects for 6 weeks. The microtomography images at 6 weeks show that the implants did not move, and untreated control defects remain empty with no evidence of a spontaneous fusion. Histological study reveals that there is newly formed bone growing up to and around the implants. There is no evidence of a fibrosis tissue ring around the implants, also indicating that there is no loosening of the implants. In contrast, the untreated controls remain empty with some evidence of a fibrosis ring around the defect hole. These results indicate good biocompatibility and bioactivity of the converted shell implants.

The condition of the periwinkle Littorina littorea, expressed in terms of its shell morphology, reproductive impairment (i.e. female sterility/intersex, male penis shedding), trematode infestation load, lipid reserves and dry/wet weight ratio, was determined in function of environmental stress along the polluted Western and relatively clean Eastern Scheldt estuary (The Netherlands). The upstream increasing pollution and decreasing salinity levels along the Western Scheldt estuary (Fig. 1) are reflected in the dry/wet weight ratio and lipid content of the periwinkles. Compared to the Eastern Scheldt, female intersex (i.e. indicator of TBT pollution) and sterility occurred more frequently in the Western Scheldt estuary, while male penis shedding was even restricted to the latter estuary. The highest population intersex and sterility incidence was found near the harbour of Vlissingen and reflects potential nautical activities. The number of trematode infested periwinkles did not differ between both estuaries, although local sampling site differences were detected within each estuary, reflecting the complex interactions that exist among parasites, hosts and the local environment. Finally, both estuaries were maximally discriminated from each other based on the shell weight of the periwinkles using a canonical discriminant analysis. Periwinkles with the heaviest shells were found in the Western Scheldt estuary and may reflect growth rate or structural population differences caused by the less favourable living conditions in the Western Scheldt estuary.

Endocrine disruption in marine populations is a growing concern. One of the best documented examples is imposex, the development of male characteristics in female gastropods, including the development of a penis and/or vas deferens, by exposure to organotins, mainly tributyltin. This syndrome has been an effective and widely used biomarker of organic tin contamination for almost 20 years. However, established imposex evaluation protocols sacrifice all the animals and repeated assessments can result in depletion of populations. To preserve animal lives as much as possible and to have a fast screening of areas subjected to organotin impacts, a non-destructive approach was tested. This approach uses reversible narcotization and a statistical criterion to evaluate the extent of imposex in the populations, instead of imposex indexes, which are based on evaluation of sacrificed, fixed animals. The data consist of two categories: males plus masculinized females in one category and normal females in another. The probability is then calculated that this sample could have been drawn from a normal population. This new method was tested in two contrasting areas of Brazil and with two different species, and the results compared to the conventional evaluation. Narcotization was completely reversed with a survival rate of 98%. Impacted sites could be identified with this technique in the same way as with the conventional one. The only limitation was that female sterility could not be verified. This disadvantage could well be compensated by the minimal mortality in the studied populations.

During the past three decades, North Atlantic populations of dogwhelks have been severely reduced in numbers, due to imposex and female sterility caused by TBT. We examined the relationship between the known history of female sterility and the present genetic diversity (measured by five allozyme-loci) and phenotypic variance (measured by nine quantitative characters) in six populations of Nucella lapillus along the coast of Norway. The environment of one of the populations was severely polluted with a vas deferens sequence index (VDSI) of 4.31, resulting in 24% sterile females. The environment of four of the other five populations was moderately polluted (VDSI 3.31-3.97, 0-1.6% sterile females), while the last population was unpolluted (VDSI 0.02, 0% sterile females). The six populations did not differ significantly with respect to genetic diversity. However, the population from the most polluted area had greater phenotypic variation in shell size, compared to all other populations. The results indicate that although TBT has had severe local negative effects on N. lapillus populations, it has not caused a reduction in long-term adaptive potential.