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Marine Pollution Bulletin 103 (2016) 5–14

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Marine Pollution Bulletin

journal homepage: www.elsevier.com/locate/marpolbul

Review

Bioremediation of waste under ocean acidification: Reviewing the role of
Mytilus edulis
Stefanie Broszeit ⁎, Caroline Hattam, Nicola Beaumont
Plymouth Marine Laboratory, Prospect Place, The Hoe, Plymouth PL1 3DH, UK

a r t i c l e i n f o a b s t r a c t

Article history: Waste bioremediation is a key regulating ecosystem service, removing wastes from ecosystems through storage,
Received 25 September 2015 burial and recycling. The bivalve Mytilus edulis is an important contributor to this service, and is used in managing
Received in revised form 17 December 2015 eutrophic waters. Studies show that they are affected by changes in pH due to ocean acidification, reducing their
Accepted 22 December 2015
growth. This is forecasted to lead to reductions in M. edulis biomass of up to 50% by 2100. Growth reduction will
Available online 8 January 2016
negatively affect the filtering capacity of each individual, potentially leading to a decrease in bioremediation of
Keywords:
waste. This paper critically reviews the current state of knowledge of bioremediation of waste carried out by
Bioremediation M. edulis, and the current knowledge of the resultant effect of ocean acidification on this key service. We show
Mytilus edulis that the effects of ocean acidification on waste bioremediation could be a major issue and pave the way for
Ocean acidification empirical studies of the topic.
Waste © 2016 Elsevier Ltd. All rights reserved.
Experiments
Ecosystem service

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
6
2. How do filter feeding bivalves M. edulis contribute to BW? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
2.1. Defining and assessing filter feeding in M. edulis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
2.2. Primary influences on filter feeding rates of M. edulis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
2.3. The role of M. edulis in BW . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
2.4. Types of waste that M. edulis bioremediate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
8
2.4.1. Nutrients, phytoplankton and organic matter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2.4.2. Toxic products of phytoplankton . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2.4.3. Examples of derivatives of burnt fossil fuel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2.4.4. Metals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2.4.5. Microplastics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2.4.6. Nanoparticles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2.4.7. Drugs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2.5. Use of M. edulis in the management of water quality. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
3. What are the key effects of OA on M. edulis?. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
3.1. Evidence of effects of OA on physiological processes related to filtration in M. edulis. . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
3.2. Effect of OA on phytoplankton. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
4. How does OA affect BW of M. edulis? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
5. Discussion and conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
5.1. OA and other stressors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
5.2. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

Abbreviations: AE, absorption efficiency; BW, bioremediation of waste; CR, clearance rate; OA, ocean acidification.
⁎ Corresponding author.
E-mail address: stbr@pml.ac.uk (S. Broszeit).

http://dx.doi.org/10.1016/j.marpolbul.2015.12.040
0025-326X/© 2016 Elsevier Ltd. All rights reserved.

2004). This shows that they can play a substantial role in the bio- turbid waters (Burkholder and Shumway. Section 2 defines how M. This in turn will reduce their to anthropogenic activities such as the burning of fossil fuel. using the car- 2002). the food source... process in BW. 2004). 2014). 2008). edulis? et al. edulis was 197 831 tons with a addition. 2011. can draw wastes deep into the sediment leading to remov. 2011). 2005). there are no studies on the effect continued global decrease in ocean pH by the end of the 21st century that OA may have on the ecosystem service of BW. Queirós et al. edulis contribute to BW? leads to reduced availability of CaCO3 for marine calcifiers (Parker 2. with a final ecosystem service with positive effects on other services too habitat range from the upper shore to the shallow subtidal (Hayward (MEA. 2013). Introduction Calcifying species play key roles in ecosystem functions (Barry et al. Melzner et al. 2013. Layman income increase. their ability to bioremediate waste. Parker et al. al of wastes by burial (Volkenborn et al. Broszeit et al. have been shown (Kroeker et al. How does OA affect BW of M. They transform the filtered material into somatic and repro. 2014).. This would be in addition Many filter feeding bivalves are vulnerable to changes in the marine to the impacts felt from temperature changes. phytoplankton. for example dominating by creating conditions for healthy fisheries and aquaculture products. For impacts on mollusc aquaculture.. cement capacity to act for bioremediation with further impacts on ecological production and deforestation.. 2007). reduced growth in M. BW supports the services of food provision and Ryland. and recreation and amenity through its contribution to bathing water M. equivalent to a 26% increase in acidity (Aze et al. The effects of OA on their primary calcifying algae and animals (Kroeker et al. 2007) and this concept has become key to linking economic explained and therefore predicting the effect of OA on these services and ecological sciences in support of sustainable environmental difficult (Cooley et al. While the economic impacts of (Norkko and Shumway.6 S. For example. edulis under OA are summarised. This 1. . Since the begin. Queirós et al. Hilmi et al.. In In 2013 the global production of M.. edulis under OA scenarios polychaetes. predicted low pH scenarios has been widely studied and reductions in Bioturbators and bioirrigators. this review and beyond (IPCC. Parker et al. Irving and Connell.1. 2005).305 (FAO. edulis filter feed. sessile assemblages on off-shore structures (Krone et al. 2015). As calcifiers.. 2011. 2013). marine microbes occur in all habitats. (2013) noted a strong impact of ocean and Larsen. It results in cleaner and less turbid water. A meta-analysis of the effects of a pH reduction by 0.. Ocean acid. ocean carbonate chemistry. Findings for molluscs (drawn addresses the third research question using examples of modelling mostly from studies on bivalves) indicate that they are particularly studies carried out on M... For fauna. known as summer of 2003 a heatwave in French waters led to massive M. remediation of waste (Lindahl et al. 1995). Such events. They are (Beaumont et al. regime used and the discount rate applied)... To our knowledge. 2013. little work focuses on the effect of OA on the filtration System Models calculated for the IPCC 5th Synthesis report project a capacity of bivalves. most living organisms can sequester wastes into their tissues value of US$ 434. Cooley et al. the movement of nutrients between demand and US$100 billion if demand increases in line with future the sediment and overlying water (Ward and Shumway. A reduction of pH also leads to changes in was timely. Thomsen et al. a common in the Atlantic from the Arctic to the Mediterranean.. Similarly. They can also be abundant. 2011. coupled with low ocean pH may ification is caused by rising atmospheric carbon dioxide (CO2) levels due lead to reduced M. reducing the carbonate ions (CO23 −) and This study aims to answer the following research questions: lowering the calcium carbonate (CaCO3) saturation of seawater. pacity to continue calcification and maintain their shells intact under degrading organic detritus and recycling nutrients (Munn. and can profoundly influence pelagic and benthic processes as well as (2012) estimate global annual losses of US$6 billion under constant add to benthic–pelagic coupling. edulis. Their ca- involved in BW. growth and mean reduction of calcification.. cascad- management (Fisher et al. growth. reducing atmospheric CO2 concentrations but at the While both the filtration capacity of filter feeding bivalves and the same time decreasing the pH of ocean surface waters. 2014). surface waters. edulis which are likely to reduce abundance for ten taxonomic groups including calcifying and non.. 2013).. through Ecosystem services are ecological components directly or indirectly the provision of habitat or by their contribution to waste remediation. 2013). Kroeker et al. Hence. Section 4 meta-analysis compared phyla only. 2011). Bioremediation of waste (BW) is an ing changes may result in the services that they provide and this even important regulating ecosystem service and can be defined as removal before extinction occurs (Barry et al. deeper light penetration due to clearer water allows filter feeders that they are used to manage eutrophic waters (Lindahl marine benthic flora to sequester carbon up to a greater depth than in et al.. Carbon dioxide dissolves into ocean functioning and wider ecosystem service delivery. changes badly affected. edulis as important trophic mode in many marine invertebrates and a key a consequence of OA can be assumed to have socio-economic impacts.. 2007. edulis. Narita et al.. Thomsen et al. How do filter feeding bivalves M. All Earth Aze et al. 2013. 2013... 2011. Filter-feeding molluscs are often found in dense populations aquaculture farmers. 2013). of waste from the environment through storage.. followed by examples of 2014). The paper is structured around these three research questions. For example. 2015).. If such species are affected by OA. edulis have been ning of global industrialisation the pH of the oceans has decreased by extensively studied (e.5 showed wastes and how they are bioremediated by M. In the Discussion (Section 5). suggested that this may particularly affect artisanal fishermen and 2008). consumed or enjoyed to produce human well-being (Boyd and The role of species vulnerable to OA in these services is not yet fully Banzhaf. Hüning et al. 2004. mollusc harvests between 2007 and 2060 (according to the future CO2 thos through faeces and pseudofaeces (Ward and Shumway. Filter feeding is an ocean acidification are not well studied. 1995). 2013). such as burrowing shrimps or several key physiological functions of M. Filter feeders actively pump large volumes of water For example. in the environment particularly a reduction of ocean water pH. M. in a review of the potential impacts of OA on Mediterra- over a filter that collects highly dilute material for feeding (Riisgard nean countries. For example.. These changes to ocean carbonate chemistry and pH have 3. Effects include significant reductions in adult and larval to management options as well as human health implications of eating survival. In this way they improve water quality by removing acidification on Mytilus species (edulis and galloprovincialis) and suspended particles (seston) from the water column (Grizzle et al. 2009). are also briefly discussed. (2009) estimate an annual et al.. they are also In the marine environment many animal taxa and guilds are known to be vulnerable to changes in OA (Kroeker et al. 2013. They may provide services to other species. calcification. 2013). edulis form an interesting case study because they are such effective quality. Section 3 summa- negative effects on survival. spat die-off (FAO. development and rises research into effects of OA on M. for example. edulis production. 0.. 2013. Aze et al..g. 2013). effect of OA on calcifying organisms such as M. edulis ocean acidification (Kroeker et al. loss to the US of US$75–187 million of direct revenue from decreasing ductive growth and aid the deposition of particulate matter to the ben. edulis? large effects on marine animals which have been the focus of sustained research effort in recent years (Melzner et al. 2005). bonate ions from seawater to form protective shells. burial and recycling This research focuses on the bivalve mollusc Mytilus edulis. 2014).. / Marine Pollution Bulletin 103 (2016) 5–14 1. What are the key effects of OA on M. Also.

Krone et al.03) − Field Hawkins et al. edulis is beneficial to understanding their capacity for BW. (2011) Filtration rate (FR) Amount of water Not given mL min−1 33. One way to measure filtration capacity is to mea- cation as well as sediment.004) 3(1/OCI)] matter that is absorbed during passage through the digestive system . particular. TPM = total particulate matter. Clearance rate. 2013). it is important to know the volume of water that has been recirculated as it reduces the efficiency of 2. Bivalves.. sessile lifestyle and because they can dominate hard sub. uals or themselves. are often used for contaminant monitoring due to their necessary (Møhlenberg and Riisgård. Measure of filtration Definition Calculation given Unit used Result Setting Reference (as given in the paper) Filtration capacity Amount of water filtered in Not given mL min−1 33–50 Field Lindahl et al. They makes it difficult to compare measurements from different studies..1. OC = organic carbon. particularly for application in coastal management measures. reducing effects of eutrophi.5–6 Lab MacDonald et al.. (2010) ingested dietary organic matter that is absorbed during passage through the digestive system Assimilation efficiency Percentage of total Not given % 1. The literature provides a range of ing on the question addressed in a particular study. edulis to filter large volumes of unfiltered water.. 2011). This measure does not incorporate study the fate of persistent organic pollutants (McEneff et al. they have been used to feeding ratio (Riisgård et al..91(±0. Defining and assessing filter feeding in M. (1998) are coefficients not further explained in the original manuscript. 2013). edulis can also adjust it by closing their valves when ly mussels.28) ∗ TPM ∗ Field Hawkins et al. Riisgård et al.36 Lab Bricelj and Kuenstner (1989) matter taken up from the water column Assimilation efficiency Not defined Not given % 24–38 Lab MacDonald et al. and assimilation efficiency (sometimes To understand the role that M. edulis M. edulis plays in the delivery of called absorption efficiency. provide important secondary habitat on hard substrata. 2011). (2011) siphon −1 Clearance rate Not defined Not given mL min 33 Lab MacDonald et al.43) Assimilation efficiency Percentage of organic Not given % 91. (Lindahl et al. 2014) recirculation of water that has already been taken up by other individ- and metal pollution (Chase et al. (2011) Clearance rate Volume of water filtered Not given Lh−1 g−1 2–12. b.13 ∗ (±9. showed that they can cover the structures with up to 3. Broszeit et al. 1978. filtration rate.. Clearance rate (CR) measures for bivalve filtration but they are not clearly defined or is a common indicator of M. filtration rates. removing large quantities through an assemblage of filter feeders in a set amount of time of phytoplankton and therefore nutrients.34) ∗ OC2... 2011. / Marine Pollution Bulletin 103 (2016) 5–14 7 2. depending on cell Lab Bricelj and Kuenstner (1989) completely free of particles abundance per unit time Filtration rate (FR) Volume of water cleared of Not given Lh−1 FR dependent on food Lab Widdows (1978) particles per unit time concentration. The most basic parameter to describe filtration physiology in measurements of M. and and units. While this diversity of filtration parameters in M. m−2 (Krone et al. edulis biomass on offshore wind energy structures M.. (1998) 1. (2011) Filtration rate (FR) Not defined FR = a ∗ TPMb ∗ OCc mg h−1 4.6 Lab Melzner et al. They reduce the size of the gape when phyto- strata both in terms of weight and abundance compared to other sessile plankton cell concentrations are too high or too low for their optimal species (Widdows et al.1–41 Lab Riisgård and Møhlenberg (1979) transported through the gills = pumping rate Filtration rate (FR) Not defined Not given mL min−1 9.4 kg of biomass is filtration or pumping capacity (from now on filtration capacity). However.64–92. For example. How do filter feeding bivalves M. (2010) ingested dietary organic matter that is absorbed during passage through the digestive system Assimilation efficiency Percentage of total Not given % 81–90 Lab Reid et al. OCI = organic carbon ingested. They often surements and it highlights the inconsistencies as concerns definitions dominate fouling communities in the shallow subtidal as well. 2005). 1982. animal and M. size of animal and temperature Filtration rate (FR) Not defined Not given mg h−1 2. it also Mussels of the genus Mytilus occur worldwide on many coasts. (2011) Assimilation efficiency Percentage of total Not given % 54 Field Reid et al. dominate hard substratum communities and have a well-developed Table 1 lists definitions used by different authors as well as units of mea- and efficient filtering system (Brzozowska et al. edulis feeding activity and measures the Table 1 Different types of filtration measurements taken from the literature. from now on assimilation efficiency) are bioremediation of waste it is first necessary to understand how filtration also used to describe filter feeding efficiency and are measured depend- is documented in the literature.15 ∗ (±0. 1995). a. S. 2001).149(± 0. (1998) ingested dietary organic [0.. harmful bacteria and contaminants sure the size of the exhalant siphon as this is controlled by the size of the (Birkbeck and McHenery. edulis. They can lead to ecosystem changes because This measures the amount of water going through a filter feeder or of their filtration capacity (Krone et al. For example.26 ∗ (±1. (2005) a given time Exhalant siphon area Size of open exhalant Not given mm2 16–49 Field MacDonald et al. 2012). MacDonald et al. edulis contribute to BW? consistently used. 2013).. and c in the calculation of FR in Hawkins et al.

Toxic Derivatives of Metals Microplastics Nanoparticles Drugs number phytoplankton phytoplankton burnt fossil (figure 1) and organic matter fuels Cycling Growth ✓ Cycling Detoxification Not always ✓ Sequestration Bioaccumulation ✓ ✓ ✓ ✓ ✓ ✓ Export Excretion through faeces. through three mechanisms (Table 2): firstly through as it gives the volume of water (or time spent clearing water) that is cycling/detoxification. 2004). 1979) measure FR as a volume per unit time and their tissues. and metabolise them of water cleared of particles per unit time and this definition is similar to a less toxic form (example below) (Baumard et al.. This reduces the damaging than just the total amount of water passing through the individual. some toxins become more toxic rather than being detoxified. Both types of faeces 1996). get buried in sediments.. edulis Melzner et al. assimilate the particles. have a higher mass of particles than small particles of seston. This way. to the definition of CR given by MacDonald et al.. measured in the outflow of a control chamber that contains no animal (MacDonald et al. benthic and lateral export. Ciliary (13%) and 5–21% amino-N via urine. 2011). edulis die or are ripped off their viscosity. However. due to hydrodynamic processes. toxic hydrocarbons act on indirectly through the digestive system when the compounds are solid Table 2 M. Rather than measuring the mass of seston 2. edulis column. edulis from the Baltic Sea Waste can be defined as “materials for which there is no this decline of ciliary activity due to low temperatures led to a reduction immediate use and that may be discharged into the environment” in feeding rates of 35% (temperature difference approximately 8 °C) (Hinga et al. 2011. Larsen and Riisgård production (Burkholder and Shumway. For example. Faeces are materials For this measure. 2011). 2004).. edulis participate in BW through sequestration and subsequent capacity defined by Lindahl et al. 2009). (2011) or the filtration M.. 1998).4. Thirdly. they how much seston that volume of water contained. The role of M. edulis bioremediate ciliary activity in M. edulis have been undertaken in laborato. taminants stored in their tissues are also moved to the seafloor and bur- reduced temperature also increases viscosity of the seawater which ied. M. They use processes that sequester waste in such a way that it (1979) clarify that when there is no recirculation of water within is no longer biologically available in the water column and does not M. edulis is influenced by water temperature and water periods of time (Newell.1 and 2. from the seston mass 1995). in this case. important in benthic–pelagic cycling and burial. There are several definitions in the literature causing filtration rate they can take up toxic wastes from incomplete combustion of fossil (FR) to be an unclear term. These materials are stuck together by mucus during and is measured by comparing organic matter in the faeces to the organ.8 S. rather than further underlying biological rea- sons such as reduced metabolic rates. they fall to the seafloor and. For example. they excrete nitrogen in the form of NH+ 4 (70%). Hawkins et al. Types of waste that M. edulis in BW lost per time (for example by weighing filtered seston from the control chamber) it is often calculated as volume per time without indication of Once seston has been filtered from the water by M. varying by waste and process. (2009) suggest after careful evaluation of the literature that increased viscosity due to lower temperature is solely responsible for reduced 2. This can change the way organic matter is then transported through the water 2. toxicity does occur when M. They (Riisgård exhibit toxicity. edulis.. CR is more infor. While temperature affects metabolic rates (Widdows. this is done them as narcotics leading to a depressed clearance rate and diminished by subtracting seston mass remaining in the outflow of a treatment scope for growth through loss of feeding opportunity (Widdows et al. metabolic rate and the size of the individual mussels (Riisgård et al. They pro- measurements from different areas and studies (Table 1). edulis can take up wastes via two pathways: di- (Melzner et al. as described in Sections 2. this includes atmospheric. 1978). For M. (2005).. In experiments.. con- 2011). This excreted nitrogen is bioavail- activity is the movement of specialised cell organs within gills that able and can lead to renewed phytoplankton and microphytobenthos create a water current allowing bivalves to feed. Widdows (1978) defined FR as the volume fuels such as polycyclic aromatic hydrocarbons. pseudo-faeces ✓ ✓ ✓ ✓ ✓ Export Excretion through urine ✓ ? ✓ Not always: during metabolisation. Therefore they et al. The majority of studies a collection of materials that are either selected because they are not carried out on filtration in M. (1998) and are consumed by other species including humans (Mebs. edulis also use mucus to laboratory and field measurements have been found (Hawkins et al. 2015). the type and the availability of food in the water column. This makes it difficult to compare system. the support by strong wind and wave action. Above a certain threshold of food (cells ml−1) both types may not be very meaningful in the field. M. Additionally. Newell. Contaminants may also influence the filter rect absorption of the compound in the water phase through the gills or feeding rates of M. and disagreements between of faeces can be produced simultaneously. chamber containing an individual of a M. edulis. Once M. 2011). If it is dense it may sink faster but if it is less dense it may remain in the water column and be available to other species for longer Filtration in M. FR is equal to CR.2. Broszeit et al. . often using single species of algal cells as food. the passage through the digestive system. MacDonald et al. bind pseudofaeces together (Riisgård et al.. that have passed through the digestive system from where nutrition AE is the percentage of organic matter taken up from the water column has been extracted. urea reduces the rate of ciliary action (Larsen and Riisgård. edulis or in a laboratory aquarium.3. 2011). Still. 2011). edulis. (2011) follow suit. duce two solid filtration products: faeces and pseudofaeces which are Another variable in filter feeding is the assimilation efficiency (AE). Riisgård and Møhlenberg storage.2.. 1999). Process Mechanism in mussel Nutrients. and hence mative than filtration capacity with regard to bioremediation of waste participate in BW. (2011) measure FR as the amount of seston removed by aiding export through all the processes that transport wastes out of a and display it as a weight per hour. the definitions are most similar across publications. They use metabolic processes that change wastes cleaned of seston after going through an individual per unit time rather into harmless or less toxic compounds. food or because there is too much food in the water column (Riisgård ries. edulis. / Marine Pollution Bulletin 103 (2016) 5–14 amount of seston removed from the water. edulis contribute to BW in several ways. effects of such wastes on themselves and other species. Pseudofaeces are made up of ic matter in the diet (MacDonald et al. for example by storing toxins from phytoplankton in and Møhlenberg. Secondly. Primary influences on filter feeding rates of M.

Diaz. In M. edulis.. Heip. This means that M.. 2001). Diaz and Rosenberg. for medicine. 2014). lead.2. edulis isms (Thompson et al. they can cross into the hemolymph.4. A study by Tedesco et al. 2013). accumulating substantial amounts of some of these toxins because they are not affected by them (Moroño et al. Rabalais et al. 2001). edulis. digestion and effects on organisms. Brzozowska et al. 1999). can grow because of the nutrients in the water column. bioavailability and uptake.. Cole et al. The 2. can have positive effects on metals than larger ones.4. For example. Such contaminants can 2008). in the gills and none in the mantle tissue. edulis in the management of water quality (Samanta et al.. Broszeit et al.. edulis in the Baltic Sea revealed that M. 1995. These authors also showed that ex- Excess nutrient loading (eutrophication) due to an imbalance in the posure to microplastics also increased energy consumption by 25% nitrogen cycle caused by. 2001.4. They also Sea. This work was to BW. cosmetics and technical equipment. 2011). (2009) calculated the cost- remove heavy metals from seawater. edulis accumulate PAHs in their tissues Lindahl et al. Microplastics leads to increased growth of phytoplankton and greening of the water may also transport contaminants into exposed organisms as these accu- column (Riebesell. Phytoplankton and organic matter are primary food sources of Depending on size. edulis can readily accumulate lipophilic organic compounds. A study carried out on concentrations of PAHs A number of studies have investigated the role of M. edulis bioremediates these at current occurring in the pelagic zone as well as in sediments and marine organ- CO2 levels are discussed in turn here. edulis accumulated in the digestive gland. 2004. edulis can take them up via feeding (Thompson et al. Gren et al. ability to select metals they can take up as an important mechanism Models to assess carrying capacity of coastal ecosystems for M. They are highly toxic. 2012). consequences. 2010). hence representative Microplastics (b1 mm) are ubiquitous in the marine environment examples of wastes and how M. edulis (Baumard et al. edulis to ensure that enough trace metals are taken in for their metabolism and other types of fish and shellfish aquaculture are well developed and (Brzozowska et al. and ecosystem service. Examples of derivatives of burnt fossil fuel bioavailable and can be taken up by M. particularly if M. edulis are important in reduc. one (O'Driscoll et al.6. 1989.3. In the water column. plastics from the environment or if they egest them. edulis which they then convert into biomass (Riisgård et al. nickel and chromium) in two size of using the organic carbon of the salmon faeces as well as excess classes of Mytilus sp. / Marine Pollution Bulletin 103 (2016) 5–14 9 (Baumard et al.4. edulis can either remove in shallow bays and enclosed seas (Diaz and Rosenberg.. S.4. Dying size they end up in waterways and ultimately in the marine environ- and dead phytoplankton is digested by microbes reducing dissolved ment. edulis. Their results indicate that this aquaculture. where they may be stored long-term... However. edulis in the management of this pollution... mulate onto the particles (Mato et al. (2012) measured the and found that if they are placed in the actual plume they are capable uptake of heavy metals (zinc. these par- 2. remove nanoparticles from the system by accumulation. phytoplankton and organic matter ticles are either egested in faeces or remain within the individual.1.4. and then either accumu- Polycyclic aromatic hydrocarbons (PAHs) are products of fossil fuel late in their tissues or become metabolised (Celiz et al. Reid et al. 2009).7. edulis in faeces plumes of salmon cages In a short experiment (24 h).4. M. Nutrients. Metals assimilation efficiency of M... 2011). 2008. Their impacts on marine deal with each of the waste types are also summarised in Table 2. or be stored in M. Therefore.. For example.5. edulis do the less dangerous benzo[e]pyrene (B[e]P) which was shown in the not feed on nutrients directly but on the phytoplankton biomass that ratio of B[a]P to B[e]P within the tissues of M. 2002). after digestion. study carried out in Ireland found 80 pharmaceuticals and their metab- olites in municipal sewage effluent (McEneff et al. lation of organic matter in the form of living and dead phytoplankton has far reaching ecosystem.. hydrodynamic . Drugs these compounds into less harmful products which they then egest Pharmaceuticals and their metabolites occur in coastal waters.. Broszeit et al. meaning they found less of a effectiveness of using M... Yet. 2005). leading to their increased example toxins produced by phytoplankton.5. Studies so far show that nanoparticles can cross and damage biological membranes and cause oxidative stress in metazoan cells. 2009. They are capable of abundance in the marine environment. for example the carcinogenic benzo[a]pyrene (B[a]P) into bility of using M. leading to hypoxic zones particularly Cauwenberghe et al. primarily M. Microplastics each waste varies depending on the type of waste. and organic matter combustion.. Gooday et al. (2005) tested the feasibility of using M. edulis 2010. they are available for filter feeders such as M. (2010) measured the 2. The role of M. Nanoparticles abundance of phytoplankton in surface waters leads to a reduction of Nanoparticles are particles of size b100 nm and due to their small light penetration and hence photosynthesis in deeper waters. They are 2. edulis aquaculture and this can reduce their filter capacity as well as their reproductive as a way of reducing nitrogen waste (N) in the Eastern Skagerrak. edulis can biotransform reducing excessive nutrient loads in coastal waters and to test the feasi- some PAHs. widely used. that they will be it is likely to increase further due to sustained human population contained within faeces and therefore more likely to sink to the seafloor growth and resource intensification (Rabalais et al. including M. edulis in in M. edulis in the bioremediation of 2. Toxic products of phytoplankton are increasingly developed and used for a number of purposes such as M. They also transform 2. This accumu. The processes and how M. also coupled with market valuation for bivalves and evaluation of an N market is being implemented in Sweden and Norway. 2004).4. 1999). 2008). (2010) showed that gold nanoparticles oxygen in the water column which can lead to hypoxic and anoxic fed to M. edulis showed that smaller individuals are less capable of selectively absorbing can be sold for human consumption. they can show how physical.. Globally. 2014). edulis farms to abate nutrients in the Baltic reduction of chromium than the other three metals tested. Nanoparticles 2. effectively reducing their contribution Sweden and demonstrated improved water quality. river run-off from agricultural activities when compared to those not exposed to microplastic. 2015).. Their results indicated that they can selectively feed coming from the cages. carcinogenic and mutagenic to marine and terrestrial animals and humans 2. the digestive tubules and gut cavity. Coastal eutrophication is one of the biggest threats to marine then be moved through the food chain to higher trophic levels (Van ecosystems and their functioning. ecosystems are still poorly understood but it has been demonstrated that marine invertebrates. This means that M. 1995). Use of M. M. This indicates that mussels develop the nutrient levels and be economically feasible too. a smaller portion zones (Diaz and Rosenberg. success (Eertman et al. little is ing phytoplankton biomass and organic matter and thereby the known about the effects of nanoparticles on the environment or their negative effects of eutrophication on the marine environment. following a model by the US (Lindahl et al.

negatively affected by a reduction in ocean pH of 0. edulis grow: effectiveness of M. growth under current pH conditions (pH 8. 7. however. 2005). This section therefore concentrates on food is available (Melzner et al. They often do not take adaptation and evolution. 2010). growth is necessary to protect the soft tissues. for example in the Late Permian Extinction. Melzner et al. immune responses or internal pH were excluded from this review as ary mechanisms into account nor biological or other interactions it can be argued that they are not directly related to filtering capacity. For the purpose coastal ecosystems for aquaculture can also be useful in assessing the of this review we concentrate on two ways in which M. 2011) and that somatic growth is the effects of OA on these physiological traits. However.. systems of the Italian island Ischia (Hall-Spencer et al. as they will their metabolic rate to maintain internal pH. a 7.1. 2014. and also because the animals were fed at higher than Navarro et al. edulis in abatement of pollution (Lindahl et al.. 2011). of the species studied. excretion of NH4. 2013).94. They found that high CO2 conditions water exchange produce less growth in M. This level).1 to 6. edulis lasted six months reduction in oxygen uptake indicating a metabolic depression.10 S. using a pH range from 8.. laboratory or field experiments. edulis Capitellidae sp.. 2010. M. Hilmi possibly due to the short time-frame of the experiment. On the other hand. 550. increased metabolic rates and suggested that the gastropods increase Overstocking of M. Field experiments might In a comprehensive meta-analysis..10) to growth in pH 7.1. edulis. Thomsen and colleagues carried out several experiments lasting between five weeks and two months (Thomsen et al.14 to (Hilmi et al. The CO2 concentration in experimental tanks can be which some are calcified (Furuhashi et al. They lasted from 20 days areas of long-term streams of CO2 and in that they are open to naturally to six months (Table 3).14 Thomsen and Melzner (2010) demonstrate that metabolic rate under (4000 μatm).. However. Most studies measured several parameters. 2009). et al. edulis in the field (for example. (2015) measured metabolic rates of farms (Waite et al.. edulis exposed to four levels of pCO2 (380. Only one experiment looked at survival. (2013) exposed juvenile Mytilus chilensis for 70 days to natural rates which may help them invest in shell growth. et al.. Filgueira et al. It lasted for 44 days and found reduced survival at 3. 2009). They also found that shell growth is suppressed from pH 7... 2005..5 and 7. edulis is influenced by the following physiological factors: metabolic 2013).01 OA conditions first increased at a pH of 7. 2010). (Harvey et al. termed the ‘Lilliput effect’ (Garilli et al. Keppel et al. Laboratory shell thickness or it was split into organic and inorganic growth as experiments may both under. million (ppm) or measured gas pressure (atm/μatm). Slower growth will therefore lower the capacity to filter and Shumway.0 over 36 h (12 h at each pH while all shell growth parameters increased under lower pH.19 Thomsen et al. The shortest experiment species sensitive to high CO2 concentrations to avoid such areas lasted 20 days and the authors used scenarios ranging from pH 8. Evidence of effects of OA on physiological processes related to filtration pH 7... edulis tissue growth in aquaculture (Wang et al. It has been shown experimentally that M. 2014). different units are shell growth occurs when several layers of shell are produced of cited in this section. in M. 380. / Marine Pollution Bulletin 103 (2016) 5–14 and biological parameters can differ within bays and how these conditions with pH of 7. other studies...7 and 7. have been carried out looking byssus thread attachment significantly deteriorated under high OA at the effect of OA on bioremediation of waste. Mytilus with M.. Kroeker et al. No experiments.3 as opposed to 8. Broszeit et al. 2015). 2011. The models used to assess carrying capacity of feed by reducing biomass at any given point in time. 2015).2 of this article.. 2013). Growth was reduced in animals .7 (1021 μatm). such as calcification. Grant et al. To allow shell growth. 2010. 700 and 1200 ppm of pCO2 with results showing a significant The longest study on OA effects in M. Gazeau et al. In an experiment lasting 35 days using pH range of 8. scenarios. Negative effects are in respiration rates unless temperature was increased for the same often localised to the aquaculture farm and can include low biological amount of time. They found that shell growth can remain stable if sufficient rate and size of the individual. Several parameters measured and displayed in several ways.. by providing in M.1 in the control differences are reflected in M. As stated in Section 2. animals must be able to calcify and this is metabolically costly under 3. Only few studies measured metabolic rate under OA in M. compete with each other for food. edulis Melzner. the bivalve Mytilus galloprovincialis.7.. and then decreased at higher to 7. pH levels (while remaining above the metabolic rate in the control However. 2008.7. edulis (Waite et al. the filtration capacity of Thomsen and Melzner. edulis? OA (Garilli et al.1 and reduced shell growth at pH 7. diversity with a prevalence of opportunistic species such as polychaetes Shell length correlates significantly with pumping rate of M. Garilli et al. somatic growth which leads to an increase in soft tissue while shell 2005.. Additionally. They found a ments of OA effects on M.5 (O'Donnell et al... 2011). Hilmi et al. 2008. two Mediterranean gastropod species (Nassarius corniculus and Cyclope Areas within a bay with low seston concentrations due to reduced neritea) near CO2 vents in Italy. is not found near natural vent shell volume growth among the nine treatment levels they used. (2015) compared from the southern Baltic Sea have a local adaptation to low pH values. Natural CO2 vents in shallow marine All studies that measured parameters affected by OA relevant to BW areas can aid research into future high CO2 environments. 2013). unaffected by low pH (Thomsen et al. (2013) showed provide more realistic scenarios than laboratory experiments but are that molluscs (the study summarised results at phylum level) are technically difficult to carry out. they found a significant decrease of inorganic shell animals). No significant changes in resting metabolic rates were found in could be due to the smaller decrease in pH treatment compared to this study.. 1992) and size of individual organisms depends on their leads to anoxia and build-up of toxic hydrogen sulphide (Burkholder growth rate. Other parameters that are OA. but occurring assemblages.. edulis were carried out in the laboratory. build-up of faecal matter which then (Jones et al. Jakubowska and Normant (2015) exposed individuals of this species After a 10 week exposure there was no effect on somatic growth to gradually reducing pH of 8. edulis can also lead to reduced growth. Thomsen et al.7 for a period of 10 months and they found no significant reduction enclosed systems such as bays (Reid et al. they only affect small areas allowing only those relevant to BW are listed in Table 3. Due to the variety of ways of expressing 2015). 2012). Therefore there are only few experi. led to smaller body sizes The effects of OA on marine organisms can be studied either by of many molluscan calcifiers. 750 and coruscus showed a significant reduction of respiration rate under OA 1000 μatm) (Fitzer et al. What are the key effects of OA on M. Gren et al. Previous OA events due to volcanic activity. below the farms. Other studies have worked with closely related species.. (2014) exposed Mytilus galloprovincialis to pH changes of ents in the form of faeces and urine may lead to negative effects on 7. 2010: Melzner et al. mass.6 (Berge et al. edulis trossulus growth at a pH of 7.and overestimate reactions of species to well as aragonite and calcite growth.. edulis. because of their relatively short duration compared to the longevity measured in OA experiments. They found no significant differences in species closely related to M. For example. (2013) found no differences in shell growth. 2010. 2006). for example as parts per for shell growth can be measured such as changes in length. Thomsen mean 17% reduction in growth in all mollusc studies they assessed. Shell growth and calcification are not interchangeable because pH changes and CO2 concentration within the studies. Thomsen and Melzner. their own input of nutri. 2013).5..

3350 μatm/8.01. 2014). 2114. 2014)..94 Keppel et al.000 ropes will produce 13% and 28% less suggest that OA may lead to a size reduction in phytoplankton. 7. Experimental Experimental treatment: CO2/pH Parameter measured Impact CO2/pH level that first caused Author(s) duration significant change 20 days 300. Adding these values together for M. 142. (2013). edulis scarce. M. 1000 μatm/NA and Shell growth ↓ 550 and 750. 750. 405 Pa/NA. 6. (2013) 7. In general.08 Somatic growth ↔ 2 months 385. 2400. edulis feed most effectively on three levels of pCO2 (380. chilensis and while the capacity to filter particles b 1 μm is reduced to 15% (Canesi found that with time. Animals under decreased pH showed reduced shell weight and fresh weight growth (Gazeau To understand the impact of OA on M. high or Shell growth low food ↓ 405 Pa Melzner et al.1. 7. plus 4 °C Whole animal wet mass ↑ 760 ppm/7. Reid et al. While growth was not affected by reduced pH alone. 7.14–7. Additionally.2. While the effect of OA on filtration parameters has not been studied With their filtration efficiency. (2013) also calculated production under OA to the small size of the alga. Shell growth ↓ 4000 μatm Thomsen and Melzner (2010) 7. Research into the effect of OA on feeding physiology of understand how OA will affect their primary food source: phytoplank. 2011. 2010).56. filter feeding depends not only on external factors eutrophication to such an extent that M. How does OA affect BW of M. (2015) measured several ton. chilensis for 70 days to ecosystem (Harvey et al.10. edulis were reduced due M. edulis. edulis contribute to BW in several ways.14 Somatic growth ↔ Metabolic rate (oxygen consumption) first ↑. particularly OA can lead to bottom-up control of the (2013) exposed the closely related species M. Additionally.7 Survival ↓ 7. Reduced growth was also found for M..94. 1120. For example..50 35 days 472. 600. / Marine Pollution Bulletin 103 (2016) 5–14 11 Table 3 Effect of OA on M.4. Evidence on the impact of OA on metabolic rate is more under OA scenarios. One study tested if metal pollution on M. galloprovincialis 3. Yet. In 750 ppm and 1200 ppm scenarios a typical Chilean aqua- how phytoplankton communities will change under OA.7. (2015) ambient temperature. 1992). 7.05. 1500 μatm/8. Bricelj and Kuenstner (1989) found that in nificant decline in CR. a management tool to clean up bays and coastal areas. Phytoplankton form the base of the marine food web and are crucial metabolic indicators under OA and increased temperatures in for biogeochemical cycling. 1400. Therefore it is important to understand scenarios. Effect of OA on phytoplankton under a 0. 750 and 1200 ppm). Shell volume growth ↔ O'Donnell et al. They measured clearance any particles with sizes N6 μm with a filtering capacity of 90%. 7. (2006) Shell growth ↓ 7. Size is one crucial factor in the ability of M.6. Shell growth ↓ 4000 ppmv Thomsen et al. they showed a sig- et al. 2005. Turley and Gattuso. also Shell length growth ↑ 760 ppm/7. because a larger individual can filter more water. edulis. Parts of the table are reproduced from Parker et al..3 pH unit decrease for 10 months. Units of CO2 and pH measurements are taken from each paper but cannot be standardised as insufficient information was provided to carry out conversion.1 Berge et al. This may subsequently it does not account for non-linear changes to these estimates) there lead to an increased likelihood of hypoxic zones (Tagliabue et al. chilensis. edulis in terms of growth impacted by OA. to filter feed (Jones et al. phytoplankton species abundance and distribution will affect M. In the same study of anophagefferens (2–3 μm) the CR and FR of M. (2014) control temperatures or 2 °C Calcite growth ↑ 1000 μatm temperature increase Aragonite growth ↓ 550 μatm exposed to 750 μatm and above 1000 μatm. This growth was compen. 500.. climate change. they show reduced growth and survival. for M. ble to study the effects of OA on all species. Some models culture farm with 10. Their enormous diversity makes it impossi.38. 240. aquaculture (Lindahl et al..7. S. 7. (2011) low food Shell growth high food ↔ 6 months 380. 7. Navarro et al. 1120 μatm then ↓ 10 weeks 400. 1100. in the highest pCO2 treatment. 18%. 1021. 2012). 2014). This change in in absorption efficiency) under the 1200 ppm scenario may occur.. 550. (2013) 1200. it is also important to et al. (2010) 7. edulis are negatively pH scenarios will have a negative effect on M. sideway arrows: no significant effect. 4000 ppmv/8. also measured that in the 1200 ppm treatment.. rate (CR) and assimilation efficiency (AE) weekly on M. 7. 800. edulis aquaculture is used as such as temperature and food availability but also on the size of the . 2014). which is expected to lead to changes in phytoplankton filtration capacity of 46% (28% reduction in biomass and 18% reduction species abundances and distribution (Shi et al. 1000. As shown in Section 2.94 (in higher temperature) 7 weeks 39. chilensis. Broszeit et al. AE was significantly higher in the a brown tide of the small phytoplankton species Aureococcus control than the higher CO2 pressures.1. for example. AE was reduced by 2014). 2010.4. are no other estimates available in the literature. Curiously. the studies are widely conclusive that OA leading to low 2011). However. Shell length growth ↔ Thomsen et al. their responses to increased temperature led to a reduction in growth. 2012).94 Total dry mass ↑ 760 ppm/7. a reduction in for example iron. chilensis biomass respectively than under current conditions. arrows down: a negative effect. pH changes the character of nutrients in the sea. edulis Though this is a rather crude method of estimating this reduction (as and ultimately the remediation of nutrients. edulis as demonstrated in experimental studies. coruscus. Navarro et al. Therefore they are not consistent within the table.03. 1300. Wang et al.94 Calcified mass ↑ 760 ppm/7. MacDonald et al.. Studies discussed in Section 3 show that M.19 and high or low food Inorganic shell growth ↓ 1021 μatm Organic shell growth ↔ 44 days NA/8. because. M. edulis? et al. 4000 μatm/8. and around fish sated for by increased protein metabolism (Fitzer et al. M. 7. the experiment was carried out in tap water mixed with calcium carbonate rather than seawater.. but not at 1000 μatm Fitzer et al. edulis under different OA scenarios changed their survival and other health parameters (Han 4..94 Soft tissue dry mass ↔ Calcified mass/soft tissue dry mass ↑ 760 ppm/7. 760 ppm/8.. They example during some seasons in the North East Atlantic (Artioli et al. they aid the removal of pollution and directly in M. mytilids is scarce.6 2 months 385. Arrows up: a positive effect.

J. Rehdanz. with M. For example. edulis makes to this service.-P. However. C. Polycyclic aromatic hydrocarbons in recent sediments and mussels (Mytilus edulis) from the Western M. if society is to continue to rely on M. IPCC. edulis are crucial.. edulis hemolymph also The service of bioremediation of waste is supported by many showed reduced antibacterial action after 90 days of exposure to OA different ecosystem processes.C.. pp. (Eds. This study has shown that M. Europe in 2003. 1978). the authors found that upon expo- contribution to these processes. Finally. edulis. Coastal eco.. Increased temperature reduces the thermal tolerance of marine European Shelf. Widdicombe.. J. Hall-Spencer. A.. 2008.. This will affect M. Y. Turley.. reducing CO2 This is of particular importance to human populations because coastal emissions and thereby slowing OA and the negative effects on ecosystems provide the majority of marine ecosystem services (Worm M. edulis to con- et al. Kavanagh. Nondal. Holt. 2015) and other bivalve species (Ivanina et al.. 2012). edulis are important contributors to tion capacity of M. bioavailability and seasonal variations. P. 2014.. Their effectiveness at removing excess nutrients and feed from that are also negatively affecting the provision of marine ecosystem aquaculture sites could be considerably diminished. coupled with a reduction in the service of subject and Steve Watson. (2008) exposed M.. edulis (and other filter feeders) but also to lower the Environment. edulis will also have negative impacts on their use in coastal manage. Res. Consequently. leading to longer residence times of filtering capacity of M.M.. if M. L. M.. Gazeau et al....H. Melzner. (Ed.. species including M. edulis (e.. Budzinski. F. (2014) exposed Baumard. P. / Marine Pollution Bulletin 103 (2016) 5–14 individual M. It may be necessary to carefully regulate harvest and seeding for thorough feedback on the manuscript. An updated synthesis as the example of M. p. Gattuso. There is also a trade-off between the services of food provision Acknowledgements and BW which may be aggravated by OA. In this study. services. edulis in terms of their physiology. R.2. Filgueira et al.. and shellfishery and beach closures. J. This service is also dependent on the sure to the pathogenic bacterium Vibrio tubiashii. et al.. J. tribute to BW. edulis for growth and filtration. K. and the Department of amount of wastes entering the marine system. R.. physiological trade-off between low pH and bacterial exposure. 2010. Broszeit et al.I. J.g. M. Biogeosciences 11.. could result in expected to impact the contribution that M. 47. then low oxygen situations occur (Diaz and Rosenberg.. G. galloprovincialis to OA and increasing temperatures and showed Baltic Sea: occurrence. F. Hansson. Matear. edulis will be vulnerable to multiple stressors in the future... J. edulis hemolymph were restored. edulis abundance such Tambutte.P. fish kills. Several studies have also found reduced (unpublished).. Wakelin. Allen. L. Brander. J. such as will have knock-on negative effects on other ecosystem processes and food provision. 2008). this study shows that such. Natural Environment Research Council (NERC) and Department for Environment... H. Kurihara.1. as the: Marine Ecosystems Research Programme. A reduction in CO2 would not only lead to a reduction in The potential reduction of BW due to negative effects of OA on the negative effects of OA but also help to slow the rise of global temper- M. C.. Bellerby. Byrne.. two additional stressors ment. particularly where water exchange is reduced such as in coastal bays. Barry. edulis (around 40–50%) as modelled by Fernandes et al. edulis to four levels of pH and 5. if sea water temperatures warm and higher mortality under OA. Fossa. Butenschön. Bibby contribute to BW locally.). S. if OA leads to a decreased 2008). R.. many with the potential to reduce the bioremediation capacity of this ic and inorganic wastes. In: Hansson...L. however. the antibacterial quantity and type of wastes that are present in the marine ecosystem functions of M. Williamson. Additional studies into the effects of OA on the drodynamic forces are reduced. particularly those Energy and Climate Change (DECC) (Grant no. tions (Joschko et al. Garrigues. OA is other filter feeding bivalves (e. it is the: UK Ocean Acidification Research Programme.. Niehoff. Gattuso. such as occurred in H. the Department for BW through M. A reduction in M. P.. Effects of ocean acidification on used the combined stressors of temperature and OA and their results in. P. Bellerby. biomass 5. J.P. 2006).. edulis. co-funded by the not only important to cut down CO2 emissions to avoid a reduction in Natural Environment Research Council (NERC). Therefore. Blackford. can have negative effects on M. It is not currently feasible to quantify the contribu.T. As tion that M. NE/H017488/1) as well resulting in eutrophication. which... S. 601–612. edulis biomass could result in less harvestable biomass of M. . edulis show reduced growth in experimental animals. Further research aiming to quantify the BW carried out services such as food provision and recreation and tourism. Food and 5. that temperature alone or temperature and pH led to 100% mortality 17–47.. Food and Rural Affairs (Defra).. 2011. this will lead to a reduction of BW as predicted. this could mean that aquaculture farms may need to be kept at smaller scales. may change under a scenario of increased OA. edulis as they prefer high oxygen concentra- biomass of M. Havenhand. Montreal. edulis making an important treatments... Discussion and conclusion showed that after 32 days there was a significant reduction of phagocyt- ic activity in the lower pH treatments. combined with similar impacts on BW due to their capacity to take up different types of wastes. tative predictions of the effect of OA on BW. capacity of M. This may indicate a in a particular place. Hauton. edulis die-off during a heatwave in France in of the impacts of ocean acidification on marine biodiversity. they participate in the bioremediation of many different types of organ. Warnau. 2009.. Dizer. as well as two anonymous referees for BW. Hattam.. S.. Extreme warming events. In addition. 1999. P. C. by M. Mar. perature. C. OA is predicted to cause negative changes to M.. Roberts. marine biodiversity and ecosystem function. P. Several studies discussed in this manuscript Barry. 99..12 S. 2003 mentioned above. this will have detrimental effects on their ability to resistance to pathogens and diseases under OA in M. Conclusions and their ability to filter feed. edulis have to the service increased occurrence of harmful algal blooms. Young. Venn. OA and other stressors Rural Affairs (DEFRA) (grant number NE/L003279/1)... S. edulis would be invaluable if the ecosystem service of BW is to systems and embayments will be particularly affected because their hy. This study was supported by human consumption to preserve the service of BW. B. Additionally. atures and the increasing spread of hypoxia. J. be better understood. edulis and may also reduce their filtration rate Aze.. H. S. other mytilid species).. Dupont. Munday.. edulis for We are grateful to Jose Fernandes for useful discussions on the human food consumption. Hansen.g. edulis would also facilitate the making of quanti- polluted water in such areas (Kemp et al. 2014. In: Hennige. Pearson. Bibby et al. L.M. T.Oxford dicate that pH is a more detrimental stressor if combined with warming University Press.. (Widdows. This study indicates that their capacity to do so key species. By implication. M.. S. OA is not an isolated pressure on the marine environment but works References in concert with other stressors particularly increased sea and air tem- Artioli. 192–209 waters than on its own. eutrophication and hypoxia (Hendriks et al. Ocean acidification.). Gibbs.. Increasing levels of OA have the potential to reduce the bioremedia. Ellis et al. M. hypoxic zones of BW by depressing the capacity of M. J. Mark. edulis. J. Heterogeneity of impacts of high CO2 on the North Western 2014). Environ. J. M. 2014). Such a reduction in water quality This will have knock-on effects for other ecosystem services.

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