Mya Arenaria Research Articles

Biological effects of gold mine tailings on the intertidal marine environment in Nova Scotia, Canada

From 1861 to the 1940s, gold was produced from 64 mining districts in Nova Scotia, where mercury amalgamation was the dominant method for the extraction of gold from ore until the 1880s. As a result, wastes (tailings) from the milling process were contaminated by mercury and were high in naturally occurring arsenic. In 2004 and 2005, sediments, water and mollusc tissues were collected from 29 sampling stations at nine former gold mining areas along the Atlantic coastline and were analysed for arsenic and mercury. The resulting data were compared with environmental quality guidelines. Samples indicated high potential risk of adverse effects in the intertidal environments of Seal Harbour, Wine Harbour and Harrigan Cove. Arsenic in Seal Harbour was bioavailable, resulting in high concentrations of arsenic in soft-shell clam tissues. Mercury concentrations in tissues were below guidelines. This paper presents results of the sampling programs and implications of these findings.

Using present-day observations to detect when anthropogenic change forces surface ocean carbonate chemistry outside preindustrial bounds

Abstract. One of the major challenges to assessing the impact of ocean acidification on marine life is detecting and interpreting long-term change in the context of natural variability. This study addresses this need through a global synthesis of monthly pH and aragonite saturation state (Ωarag) climatologies for 12 open ocean, coastal, and coral reef locations using 3-hourly moored observations of surface seawater partial pressure of CO2 and pH collected together since as early as 2010. Mooring observations suggest open ocean subtropical and subarctic sites experience present-day surface pH and Ωarag conditions outside the bounds of preindustrial variability throughout most, if not all, of the year. In general, coastal mooring sites experience more natural variability and thus, more overlap with preindustrial conditions; however, present-day Ωarag conditions surpass biologically relevant thresholds associated with ocean acidification impacts on Mytilus californianus (Ωarag < 1.8) and Crassostrea gigas (Ωarag < 2.0) larvae in the California Current Ecosystem (CCE) and Mya arenaria larvae in the Gulf of Maine (Ωarag < 1.6). At the most variable mooring locations in coastal systems of the CCE, subseasonal conditions approached Ωarag = 1. Global and regional models and data syntheses of ship-based observations tended to underestimate seasonal variability compared to mooring observations. Efforts such as this to characterize all patterns of pH and Ωarag variability and change at key locations are fundamental to assessing present-day biological impacts of ocean acidification, further improving experimental design to interrogate organism response under real-world conditions, and improving predictive models and vulnerability assessments seeking to quantify the broader impacts of ocean acidification.

Large-scale mitochondrial COI gene sequence variability reflects the complex colonization history of the invasive soft-shell clam, Mya arenaria (L.) (Bivalvia)

The aim of the study was to determine genetic diversity in the soft-shell clam Mya arenaria on a wide geographical scale using mtDNA COI gene sequences. Low levels of genetic diversity was found, which can most likely be explained by a bottleneck effect during Pleistocene glaciations and/or selection. The geographical genetic structuring of the studied populations was also very low. The star-like phylogeny of the haplotypes indicates a relatively recent, rapid population expansion following the glaciation period and repeated expansion following the founder effect(s) after the initial introduction of the soft-shell clam to Europe. North American populations are characterized by the largest number of haplotypes, including rare ones, as expected for native populations. Because of the founder effect connected with initial and repeated expansion events, European populations have significantly lower numbers of haplotypes in comparison with those of North America. We also observed subtle differentiations among populations from the North and Baltic seas. The recently founded soft-shell clam population in the Black Sea exhibited the highest genetic similarity to Baltic populations, which confirmed the hypothesis that M. arenaria was introduced to the Gulf of Odessa from the Baltic Sea. The most enigmatic results were obtained for populations from the White Sea, which were characterized by high genetic affinity with American populations.

Impacts of seawater saturation state (ΩA = 0.4–4.6) and temperature (10, 25 °C) on the dissolution kinetics of whole-shell biogenic carbonates

Anthropogenic increase of atmospheric pCO2 since the Industrial Revolution has caused seawater pH to decrease and seawater temperatures to increase—trends that are expected to continue into the foreseeable future. Myriad experimental studies have investigated the impacts of ocean acidification and warming on marine calcifiers’ ability to build protective shells and skeletons. No studies, however, have investigated the combined impacts of ocean acidification and warming on the whole-shell dissolution kinetics of biogenic carbonates. Here, we present the results of experiments designed to investigate the effects of seawater saturation state (ΩA=0.4–4.6) and temperature (10, 25°C) on gross rates of whole-shell dissolution for ten species of benthic marine calcifiers: the oyster Crassostrea virginica, the ivory barnacle Balanus eburneus, the blue mussel Mytilus edulis, the conch Strombus alatus, the tropical coral Siderastrea siderea, the temperate coral Oculina arbuscula, the hard clam Mercenaria mercenaria, the soft clam Mya arenaria, the branching bryozoan Schizoporella errata, and the coralline red alga Neogoniolithon sp. These experiments confirm that dissolution rates of whole-shell biogenic carbonates decrease with calcium carbonate (CaCO3) saturation state, increase with temperature, and vary predictably with respect to the relative solubility of the calcifiers’ polymorph mineralogy [high-Mg calcite (mol% Mg>4)≥aragonite>low-Mg calcite (mol% Mg<4)], consistent with prior studies on sedimentary and inorganic carbonates. Furthermore, the severity of the temperature effects on gross dissolution rates also varied with respect to carbonate polymorph solubility, with warming (10–25°C) exerting the greatest effect on biogenic high-Mg calcite, an intermediate effect on biogenic aragonite, and the least effect on biogenic low-Mg calcite. These results indicate that both ocean acidification and warming will lead to increased dissolution of biogenic carbonates in future oceans, with shells/skeletons composed of the more soluble polymorphs of CaCO3 being the most vulnerable to these stressors. The effects of saturation state and temperature on gross shell dissolution rate were modeled with an exponential asymptotic function (y=B0–B2·eB1Ω) that appeals to the general Arrhenius-derived rate equation for mineral dissolution [r=(C·e-Ea/RT)(1−Ω)n]. Although the dissolution curves for the investigated biogenic CaCO3 exhibited exponential asymptotic trends similar to those of inorganic CaCO3, the observation that gross dissolution of whole-shell biogenic CaCO3 occurred (albeit at lower rates) even in treatments that were oversaturated (Ω>1) with respect to both aragonite and calcite reveals fundamental differences between the dissolution kinetics of whole-shell biogenic CaCO3 and inorganic CaCO3. Thus, applying stoichiometric solubility products derived for inorganic CaCO3 to model gross dissolution of biogenic carbonates may substantially underestimate the impacts of ocean acidification on net calcification (gross calcification minus gross dissolution) of systems ranging in scale from individual organisms to entire ecosystems (e.g., net ecosystem calcification). Finally, these experiments permit rough estimation of the impact of CO2-induced ocean acidification on the gross calcification rates of various marine calcifiers, calculated as the difference between net calcification rates derived empirically in prior studies and gross dissolution rates derived from the present study. Organisms’ gross calcification responses to acidification were generally less severe than their net calcification response patterns, with aragonite mollusks (bivalves, gastropods) exhibiting the most negative gross calcification response to acidification, and photosynthesizing organisms, including corals and coralline red algae, exhibiting relative resilience.

Assessment of sodium channel mutations in Makah tribal members of the U.S. Pacific Northwest as a potential mechanism of resistance to paralytic shellfish poisoning

The Makah Tribe of Neah Bay, Washington, has historically relied on the subsistence harvest of coastal seafood, including shellfish, which remains an important cultural and ceremonial resource. Tribal legend describes visitors from other tribes that died from eating shellfish collected on Makah lands. These deaths were believed to be caused by paralytic shellfish poisoning, a human illness caused by ingestion of shellfish contaminated with saxitoxins, which are produced by toxin-producing marine dinoflagellates on which the shellfish feed. These paralytic shellfish toxins include saxitoxin, a potent Na+ channel antagonist that binds to the pore region of voltage gated Na+ channels. Amino acid mutations in the Na+ channel pore have been demonstrated to confer resistance to saxitoxin in softshell clam populations exposed to paralytic shellfish toxins present in their environment. Because of the notion of resistance to paralytic shellfish toxins, the study aimed to determine if a resistance strategy was possible in humans with historical exposure to toxins in shellfish. We collected, extracted and purified DNA from buccal swabs of 83 volunteer Makah tribal members and sequenced the skeletal muscle Na+ channel (Nav1.4) at nine loci to characterize potential mutations in the relevant saxitoxin binding regions. No mutations of these specific regions were identified after comparison to a reference sequence. This study suggests that any resistance of Makah tribal members to saxitoxin, if present, is not a function of Nav1.4 modification, but may be due to mutations in neuronal or cardiac sodium channels, or some other mechanism unrelated to sodium channel function.

Measuring the impact of pollution closures on commercial shellfish harvest: The case of soft-shell clams in Machias Bay, Maine

Temporary closures of polluted coastal waters to shellfish harvesting protect human health but also generate broad socioeconomic impacts on rural, fishing-dependent communities. Improved understanding of these impacts could help coastal managers prioritize investments to protect water quality and mitigate the effects of coastal pollution. Using a regression model of monthly landings, we explore the impact of temporary closures on the commercial harvest of soft-shell clams (Mya arenaria) in the Machias Bay region of Maine (USA). We find that economic losses are significant and depend heavily on tidal activity, and the size, frequency and timing of closures. Over the nine-year sample period (2001–2009), temporary pollution closures contributed to the loss of $3.6 million in forgone revenue (2014 dollars), approximately 27.4% of total revenue. Closures linked to combined sewer overflows from the Machias wastewater system produce the majority of these losses ($2.0 million) with the largest occurring during the peak clamming season (May–August). Our results highlight the variability of the impacts of closures and the information burden for efficient management of shellfish areas and coastal waters. By strategically reducing pollution, managers could limit public health risks, avoid destabilizing harvesting and revenue, and bolster the resilience of fishing communities.

Widespread transmission of independent cancer lineages within multiple bivalve species.

Most cancers arise from oncogenic changes in the genomes of somatic cells, and while the cells may migrate by metastasis, they remain within that single individual. Natural transmission of cancer cells from one individual to another has been observed in two distinctive cases in mammals (Tasmanian devils1 and dogs2,3), but these are generally considered to be rare exceptions in nature. The discovery of transmissible cancer in soft-shell clams (Mya arenaria)4 suggested that this phenomenon might be more widespread. Here we analyzed disseminated neoplasia in mussels (Mytilus trossulus), cockles (Cerastoderma edule), and golden carpet shell clams (Polititapes aureus) and found that neoplasias in all three species are attributable to independent transmissible cancer lineages. In mussels and cockles, the cancer lineages are derived from their respective host species, but unexpectedly, cancer cells in P. aureus are all derived from Venerupis corrugata, a different species living in the same geographic area. No cases of disseminated neoplasia have thus far been found in V. corrugata from the same region. These findings show that transmission of cancer cells in the marine environment is common in multiple species, that it has originated many times, and that while most transmissible cancers were found spreading within the species of origin, cross-species transmission of cancer cells can occur.

Transmissible tumours under the sea

In some species, cancer cells can be directly transmitted between individuals. An analysis in shellfish now shows that some transmissible cancers can even cross the species barrier. See Letter p.705 Transmissible neoplasia is rare, but it has been found to occur in the Tasmanian devil (a marsupial), dogs, and recently in soft-shell clams. Here Stephen Goff and colleagues identify three marine bivalve species — cockles, mussels and shell clams — that also exhibit disseminated neoplasia. In shell clams, the cancer lineages are transmitted from a closely related species. The findings suggest that transmissible neoplasia is more common than was previously thought, and that cross-species transmission of cancer can occur, albeit rarely.

Are Medieval Mya arenaria (Mollusca; Bivalvia) in the Netherlands also clams before Columbus?

AbstractDuring the Pleistocene, the coastal marine bivalve mollusc Mya arenaria became extinct in northwest Europe. The species remained present in North America. Datings of Mya shells found in northern Denmark and the southern Baltic Sea suggest that repopulation of northwest European coasts already occurred before Columbus’ discovery of America (1492), possibly facilitated by Viking (Norse) settlers at Greenland and northeast North America. In this paper we report on findings of M. arenaria at five locations in the coastal landscape of the Netherlands: polders reclaimed from the Wadden Sea and the former estuaries of Oer-IJ and Old Rhine. The shells from four of these locations also date before 1492 AD.

Genetic evidence supports recolonisation by Mya arenaria of western Europe from North America

Genetic evidence supports recolonisation by Mya arenaria of western Europe from North America

Seasonal proliferation rates and the capacity to express genes involved in cell cycling and maintenance in response to seasonal and experimental food shortage in Laternula elliptica from King George Island

Melting of coastal glaciers at the West Antarctic Peninsula (WAP) causes shorter winter sea ice duration, intensified ice scouring, sediment erosion and surface freshening in summer, which alters coastal productivity and feeding conditions for the benthos. The soft shell clam Laternula elliptica is a fast growing and abundant filter feeder in coastal Antarctica and a key element for bentho-pelagic carbon recycling. Our aim was to assess the cellular growth and maintenance capacity of small and large clams during natural winter food shortage (seasonal sampling) and in response to experimental starvation exposure. We measured tissue specific proliferation rates, the expression of cell cycling genes, and the iron binding protein Le-ferritin in freshly collected specimens in spring (Nov 2008) and at the end of summer (March 2009). For the experimental approach, we focused on 14 cell cycling and metabolic genes using the same animal size groups. Mantle tissue of young bivalves was the only tissue showing accelerated proliferation in summer (1.7% of cells dividing per day in March) compared to 0.4% dividing cells in animals collected in November. In mantle, siphon and adductor muscle proliferation rates were higher in younger compared to older individuals. At transcript level, Le-cyclin D was upregulated in digestive gland of older animals collected in spring (Nov) compared to March indicating initiation of cell proliferation. Likewise, during experimental starvation Le-cyclin D expression increased in large clam digestive gland, whereas Le-cyclin D and the autophagic factor beclin1 decreased in digestive gland of smaller starved clams. The paper corroborates earlier findings of size and age dependent differences in the metabolic response and gene expression patterns in L. elliptica under energetic deprivation. Age structure of shallow water populations can potentially change due to differences in cellular response between young and old animals as environmental stress levels increase.

Growth of Mya arenaria L. at the northern edge of the range: heterogeneity of soft-shell clam growth characteristics in the White Sea

Linear growth of soft-shell clams, Mya arenaria L., was studied at six locations in the White Sea situated in the middle and low tidal zones. The main aim was to analyse the growth heterogeneity both within beds (with respect to individual growth) and between beds (with respect to group growth). Mya population was characterized by a slow growth and a long lifespan. Maximum shell size was 70–80 mm, and longevity was at least 25 years. In addition, growth rates were highly variable, statistically significant differences being present both within and between the beds. No differences were found in group growth between different tidal horizons of the same site. The slowest growth of soft-shell clams was observed at sites with the greatest content of fine particles (silt and clay) in sediments. Differences between beds in Mya group growth could reflect variability of feeding conditions, which were probably determined by some local features (e.g. hydrodynamic regime). However, the variation of the individual growth rate within beds was comparable to that of the group growth rate between studied locations. To a great extent it seemed to be determined by the characteristics of the initial period of soft-shell clam growth.

Shell matrix proteins of the clam, Mya truncata: Roles beyond shell formation through proteomic study

Mya truncata, a soft shell clam, is presented as a new model to study biomineralization through a proteomics approach. In this study, the shell and mantle tissue were analysed in order to retrieve knowledge about the secretion of shell matrix proteins (SMPs). Out of 67 and 127 shell and mantle proteins respectively, 16 were found in both shell and mantle. Bioinformatic analysis of SMP sequences for domain prediction revealed the presence of several new domains such as fucolectin tachylectin-4 pentraxin-1 (FTP), scavenger receptor, alpha-2-macroglobulin (α2 M), lipocalin and myosin tail along with previously reported SMP domains such as chitinase, carbonic anhydrase, tyrosinase, sushi, and chitin binding. Interestingly, these newly predicted domains are attributed with molecular functions other than biomineralization. These findings suggest that shells may not only act as protective armour from predatory action, but could also actively be related to other functions such as immunity. In this context, the roles of SMPs in biomineralization need to be looked in a new perspective.

Assessment and Molecular Characterization of Human Intestinal Parasites in Bivalves from Orchard Beach, NY, USA.

Bivalves have been shown to be carriers of the human intestinal parasites Cryptosporidium parvum and Toxoplasma gondii. The goal of this study is to determine the prevalence of protozoan parasites in mollusks of New York City using a polymerase chain reaction (PCR)-based assay. Four species of mollusks, Mya arenaria, Geukensia demissa, Crassostrea virginica, and Mytilis edulis, were collected from Orchard Beach, NY in the fall of 2014, totaling 159 specimens. Each individual mollusk was dissected to harvest the digestive gland, the mantle, the gills, the foot and the siphon. The tissues were assayed for the presence of Cryptosporidium parvum, Giardia lamblia, and Toxoplasma gondii DNA by using primers that target parasite-specific genes. C. parvum was found at a prevalence of 50%, 11.3%, and 1%, respectively, in Mya arenaria, G. demissa, and Mytilis edulis. C. parvum DNA was detected in all the tissues of these bivalve species, except the gills. Furthermore, G. lamblia was detected in Mya arenaria, G. demissa, Crassostrea virginica and Mytilis edulis at a prevalence of 37.5%, 4.5%, 60%, and 20.6%, respectively, while T. gondii DNA was not detected.

Ecotoxicological Evaluation of the Immunocompetence of two Bivalves Species (Mya Arenaria and Mytilus Edulis ) in the Saguenay Fjord Including a Salinity Gradient.

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Porewater acidification alters the burrowing behavior and post-settlement dispersal of juvenile soft-shell clams (Mya arenaria)

Although ocean acidification will impact marine organisms in the future, few studies have addressed the effects of sedimentary porewater acidification on benthic invertebrates. This study suggests that burrowing behavior and post-settlement dispersal of juvenile bivalves are altered by porewater acidification under present day conditions. We tested the efficacy of a novel method of stabilizing porewater pH using sediment underlain with food grade gelatin in both the lab and field, and then employed this method to test if porewater acidification could alter post-settlement clam dispersal under natural conditions. In the field, clams were exposed to a gradient of porewater acidification in manipulated (CO2 added) sediments for 24h to determine if acidification could alter dispersal patterns of juvenile clams under natural flow conditions; juvenile clam dispersal in the presence of different buffer types was also tested. In addition, juvenile clams were placed on unmanipulated, field-collected sediment cores in the lab which varied naturally with respect to acidification to test burrowing behavior in response to natural porewater pH. Gelatin stabilized porewater pH for 24–48h and its presence did not influence clam burrowing behavior. In the field, a significant negative relationship between the percent of clams dispersed and acidification was observed, while stabilizing porewater pH significantly decreased clam dispersal. In the lab, there was a significant positive relationship between the percent of clams burrowed and porewater acidification. This study suggests that porewater acidification has the capacity to alter the burrowing behavior and dispersal patterns of juvenile bivalves under natural conditions.

Effects of various bivalves on meiobenthic and nematode assemblages in shallow sandy sediments

A laboratory microcosm experiment was performed to study the effect of bivalves on meiobenthic and nematode community structures. Three bivalve species dominant in shallow coastal sediments of the southern Baltic Sea differing in terms of feeding mode, faecal deposition and sediment depth penetration (Macoma balthica, Cerastoderma glaucum and Mya arenaria) were selected as model organisms: our experiment demonstrated that although the bivalves had no overall effect on total meiobenthic densities, they variously affected a number of meiobenthic major taxa. Species buried under the sediment surface (M. balthica, M. arenaria) facilitated meiobenthos to penetrate deeper sediment layers compared to the surface-dwelling bivalves (C. glaucum). The most conspicuous response of meiobenthos and nematodes to bivalve activities was, however, observed at the sediment surface: densities of juvenile bivalves and rotifers, and the dominant nematode species (Adoncholaimus thalassophygas, Desmolaimus cf. zeelandicus and Halomonhystera disjuncta) were significantly reduced in surface sediments in Macoma microcosms compared to other treatments. It is suggested that different bivalve feeding modes (deposit-feeding vs. suspension-feeding), attributes of faecal deposition (to the sediment surface vs. to subsurface sediments) and depth of sediment penetration can significantly change meiobenthic and nematode communities in sandy sediments.

Analysis of Gene Expression in an Inbred Line of Soft-Shell Clams (Mya arenaria) Displaying Growth Heterosis: Regulation of Structural Genes and the NOD2 Pathway.

Mya arenaria is a bivalve mollusk of commercial and economic importance, currently impacted by ocean warming, acidification, and invasive species. In order to inform studies on the growth of M. arenaria, we selected and inbred a population of soft-shell clams for a fast-growth phenotype. This population displayed significantly faster growth (p < 0.0001), as measured by 35.4% greater shell size. To assess the biological basis of this growth heterosis, we characterized the complete transcriptomes of six individuals and identified differentially expressed genes by RNAseq. Pathways differentially expressed included structural gene pathways. Also differentially expressed was the nucleotide-binding oligomerization domain 2 (NOD2) receptor pathway that contributes to determination of growth, immunity, apoptosis, and proliferation. NOD2 pathway members that were upregulated included a subset of isoforms of RIPK2 (mean 3.3-fold increase in expression), ERK/MAPK14 (3.8-fold), JNK/MAPK8 (4.1-fold), and NFκB (4.08-fold). These transcriptomes will be useful resources for both the aquaculture community and researchers with an interest in mollusks and growth heterosis.

A second transmissible cancer in Tasmanian devils

Clonally transmissible cancers are somatic cell lineages that are spread between individuals via the transfer of living cancer cells. There are only three known naturally occurring transmissible cancers, and these affect dogs, soft-shell clams, and Tasmanian devils, respectively. The Tasmanian devil transmissible facial cancer was first observed in 1996, and is threatening its host species with extinction. Until now, this disease has been consistently associated with a single aneuploid cancer cell lineage that we refer to as DFT1. Here we describe a second transmissible cancer, DFT2, in five devils located in southern Tasmania in 2014 and 2015. DFT2 causes facial tumors that are grossly indistinguishable but histologically distinct from those caused by DFT1. DFT2 bears no detectable cytogenetic similarity to DFT1 and carries a Y chromosome, which contrasts with the female origin of DFT1. DFT2 shows different alleles to both its hosts and DFT1 at microsatellite, structural variant, and major histocompatibility complex (MHC) loci, confirming that it is a second cancer that can be transmitted between devils as an allogeneic, MHC-discordant graft. These findings indicate that Tasmanian devils have spawned at least two distinct transmissible cancer lineages and suggest that transmissible cancers may arise more frequently in nature than previously considered. The discovery of DFT2 presents important challenges for the conservation of Tasmanian devils and raises the possibility that this species is particularly prone to the emergence of transmissible cancers. More generally, our findings highlight the potential for cancer cells to depart from their hosts and become dangerous transmissible pathogens.

Field and laboratory transmission studies of haemic neoplasia in the soft-shell clam, Mya arenaria, from Atlantic Canada.

A two-year laboratory and field study was initiated in 2001 in response to mass mortalities associated with haemic neoplasia (HN) in 1999 in Prince Edward Island (PEI) soft-shell clams, Mya arenaria. A laboratory proximity experiment (cohabitation) and an inoculation challenge were conducted with clams and mussels (Mytilus edulis). Three field exposure experiments were also conducted, in which naive clams were held in sediment (in trays) or out of sediment (in mesh bags) at three high HN prevalence sites on PEI. There was a conversion to HN positive in clams in the proximity experiment and in clams injected with whole blood and cell-free homogenate, but not at statistically significant levels. No mussels or control clams became HN positive. There was a significant conversion to HN positive in as little as 24 and 58days after transfer with clams held out of sediment and in sediment, respectively. The laboratory and field experiments' results suggest that HN-infected clams are spreading the disease through water from infected clams to naïve individuals and via transplantation from affected to unaffected sites. Some environmental conditions (e.g. abnormally high water temperature and hypoxia-induced sea lettuce [Ulva lacteus] invasion) may make clams susceptible to infections or exacerbate the proliferation of HN.