Luc A. Comeau

Temperature adaptation in two bivalve species from different thermal habitats: energetics and remodelling of membrane lipids

SUMMARY We compared lipid dynamics and the physiological responses of blue mussels Mytilus edulis, a cold-adapted species, and oysters Crassostrea virginica, a warmer-water species, during simulated overwintering and passage to spring conditions. To simulate overwintering, animals were held at 0°C, 4°C and 9°C for 3 months and then gradually brought to and maintained at 20°C for 5 weeks to simulate spring–summer conditions. Changes in lipid class and fatty acid composition were related to clearance rate and oxygen consumption. We found major differences between species in triglyceride (TAG) metabolism during overwintering. Mussels used digestive gland TAG stores for energy metabolism or reproductive processes during the winter, whereas oysters did not accumulate large TAG stores prior to overwintering. Mussel TAG contained high levels of 20:5n-3 compared to levels in oysters and in the diet. This may help to counteract the effect of low temperature by reducing the melting point of TAG and thus increasing the availability of storage fats at low temperature. Mussels seemed better able to mobilise 20:5n-3 and 18:4n-3 than other fatty acids. We also found that both bivalves underwent a major remodelling of membrane phospholipids. The unsaturation index decreased in the gills and digestive glands of both species during the early stages of warming, principally due to decreases in 22:6n-3 and 20:5n-3. In digestive glands, the unsaturation index did not increase with decreasing temperature beyond a threshold attained at 9°C whereas a perfect negative relationship was observed in gills, as predicted by homeoviscous adaptation. The presence of digestive enzymes and acids in the digestive gland microenvironment may lead to specific requirements for membrane stability. That oysters had lower metabolic rates than mussels coincides with a lower unsaturation index of their lipids, as predicted by Hulberts theory of membranes as metabolic pacemakers. Both species showed increased 20:4n-6 levels in their tissues as temperature rose, suggesting an increasing availability of this fatty acid for eicosanoid biosynthesis during stress responses. The contrast between the species in TAG dynamics and the similarity of their phospholipid remodelling emphasises the essential functional role of membrane phospholipid structure and the contrasting use of TAG by oysters and mussels during overwintering.

Byssus attachment strength of two mytilids in mono- specific and mixed-species mussel beds

The mussel Xenostrobus securis is endemic to the brackish waters of New Zealand and Australia, but has successfully invaded the inner Galician Rías of NW Spain, where it coexists with the indigenous mussel Mytilus galloprovincialis. In this laboratory study, the plasticity of the byssus attachment strength of two mytilids was compared by manipulating substratum, salinity, and bed assembly. M. galloprovincialis showed stronger byssus detachment strength than X. securis, despite lower byssus coverage. Both species responded similarly to the substratum, with substantially lower byssus strength on methacrylate, which offered the lowest surface free energy. Byssus detachment values for M. galloprovincialis were lower at lower salinity. In mixed beds, a number of mussels moved upwards, eventually colonising the upper layers of the assemblage. This behaviour increased byssus strength but only for X. securis. X. securis is adapted to a wide spectrum of abiotic conditions, a trait that may promote its dissemination within estuarine environments.

Response of two mytilids to a heatwave: the complex interplay of physiology, behaviour and ecological interactions

Different combinations of behavioural and physiological responses may play a crucial role in the ecological success of species, notably in the context of biological invasions. The invasive mussel Xenostrobus securis has successfully colonised the inner part of the Galician Rias Baixas (NW Spain), where it co-occurs with the commercially-important mussel Mytilus galloprovincialis. This study investigated the effect of a heatwave on the physiological and behavioural responses in monospecific or mixed aggregations of these species. In a mesocosm experiment, mussels were exposed to simulated tidal cycles and similar temperature conditions to those experienced in the field during a heat-wave that occurred in the summer of 2013, when field robo-mussels registered temperatures up to 44.5°C at low tide. The overall responses to stress differed markedly between the two species. In monospecific aggregations M. galloprovincialis was more vulnerable than X. securis to heat exposure during emersion. However, in mixed aggregations, the presence of the invader was associated with lower mortality in M. galloprovincialis. The greater sensitivity of M. galloprovincialis to heat exposure was reflected in a higher mortality level, greater induction of Hsp70 protein and higher rates of respiration and gaping activity, which were accompanied by a lower heart rate (bradycardia). The findings show that the invader enhanced the physiological performance of M. galloprovincialis, highlighting the importance of species interactions in regulating responses to environmental stress. Understanding the complex interactions between ecological factors and physiological and behavioural responses of closely-related species is essential for predicting the impacts of invasions in the context of future climate change.

Informing Marine Spatial Planning (MSP) with numerical modelling: A case-study on shellfish aquaculture in Malpeque Bay (Eastern Canada)

A moratorium on further bivalve leasing was established in 1999-2000 in Prince Edward Island (Canada). Recently, a marine spatial planning process was initiated explore potential mussel culture expansion in Malpeque Bay. This study focuses on the effects of a projected expansion scenario on productivity of existing leases and available suspended food resources. The aim is to provide a robust scientific assessment using available datasets and three modelling approaches ranging in complexity: (1) a connectivity analysis among culture areas; (2) a scenario analysis of organic seston dynamics based on a simplified biogeochemical model; and (3) a scenario analysis of phytoplankton dynamics based on an ecosystem model. These complementary approaches suggest (1) new leases can affect existing culture both through direct connectivity and through bay-scale effects driven by the overall increase in mussel biomass, and (2) a net reduction of phytoplankton within the bounds of its natural variation in the area.

Elevated seawater temperature, not pCO2, negatively affects post-spawning adult mussels (Mytilus edulis) under food limitation

Pre-spawning blue mussels (Mytilus edulis) appear sensitive to elevated temperature and robust to elevated pCO2; however, the effects of these stressors soon after investing energy into spawning remain unknown. Furthermore, while studies suggest that elevated pCO2 affects the byssal attachment strength of Mytilus trossulus from southern latitudes, pCO2 and temperature impacts on the byssus strength of other species at higher latitudes remain undocumented. In a 90 day laboratory experiment, we exposed post-spawning adult blue mussels (M. edulis) from Atlantic Canada to three pCO2 levels (pCO2 ~625, 1295 and 2440 μatm) at two different temperatures (16°C and 22°C) and assessed energetic reserves on Day 90, byssal attachment strength on Days 30 and 60, and condition index and mortality on Days 30, 60 and 90. Results indicated that glycogen content was negatively affected under elevated temperature, but protein, lipid, and overall energy content were unaffected. Reduced glycogen content under elevated temperature was associated with reduced condition index, reduced byssal thread attachment strength, and increased mortality; elevated pCO2 had no effects. Overall, these results suggest that the glycogen reserves of post-spawning adult M. edulis are sensitive to elevated temperature, and can result in reduced health and byssal attachment strength, leading to increased mortality. These results are similar to those reported for pre-spawning mussels and suggest that post-spawning blue mussels are tolerant to elevated pCO2 and sensitive to elevated temperature. In contrast to previous studies, however, elevated pCO2 did not affect byssus strength, suggesting that negative effects of elevated pCO2 on byssus strength are not universal.

Past, Present, and Future: Performance of Two Bivalve Species Under Changing Environmental Conditions

Globally, the production of marine bivalves has been steadily increasing over the past several decades. As the effects of human population growth are magnified, bivalves help provide food security as a source of inexpensive protein. However, as climate change alters sea surface temperatures (SST), the physiology, and thus the survival, growth, and distribution of bivalves are being altered. Challenges with managing bivalves may become more pronounced, as the uncertainty associated with climate change makes it difficult to predict future production levels. Modelling techniques, applied to both climate change and bivalve bioenergetics, can be used to predict and explore the impacts of changing ocean temperatures on bivalve physiology, and concomitantly on aquaculture production. This study coupled a previously established high resolution climate model and two dynamic energy budget models to explore the future growth and distribution of two economically and ecologically important species, the eastern oyster (Crassotrea virginica), and the blue mussel (Mytilus edulis) along the Atlantic coast of Canada. SST was extracted from the climate model and used as a forcing variable in the bioenergetic models. This approach was applied across three discreet time periods: the past (1986-1990), the present (2016-2020), and the future (2046-2050), thus permitting a comparison of bivalve performance under different temporal scenarios. Results show that the future growth is variable both spatially and interspecifically. Modelling outcomes suggest that warming ocean temperatures will cause an increase in growth rates of both species as a result of their ectothermic nature. However, as the thermal tolerance of C. virginica is higher than M. edulis, oysters will generally outperform mussels. The predicted effects of temperature on bivalve physiology also provided insight into vulnerabilities (e.g. mortality) under future SST scenarios. Such information is useful for adapting future management strategies for both farmed and wild shellfish. Although this study focused on a geographically specific area, the approach of coupling bioenergetic and climate models is valid for species and environments across the globe.

Physiological changes during overwintering of the Eastern oyster Crassostrea virginica (Gmelin, 1791)

This study aims at better understanding of the physiological changes that occur during the winter months in Eastern oysters (Crassostrea virginica) living at their northernmost distributional limit (48°N). It was conducted in northeastern New Brunswick, Canada (47°49′N, 64°46′W). Oysters were held in Vexar bags on the bottom of a bay and collected before (December) and during (January–April) the ice-cover period. Winter mortality was <5%. A significant decrease in the concentration of lipids stored in the digestive gland (from 42.0 to 22.5 mg/g wet tissue) indicated that the animals had access to oxygen, probably through minimal shell openings. Proteins were also used, although less heavily (decrease from 68.0 to 59.1 mg/g wet tissue), leading to an accumulation of ammonia in the intrapallial fluid (up to 0.83 μM), which is modest considering the length of time the animals lived as almost closed systems. In contrast, carbohydrates appear to play a minor role in winter energetics. Low pH values of intrapallial fluid were noted in late winter (c. 7.0) as compared with those measured at the onset of winter and in spring (c. 7.7). The animals used more energy substrates in late than in early winter, which might be partly related to the necessity to intensify detoxification mechanisms as toxic metabolites accumulate. Despite the fact that oysters do not filter-feed at temperatures <0 °C, the mass of the digestive gland increased by a factor of 1.5 during early winter, presumably as the result of nutrient transfer from other tissues. The ability of C. virginica to use energy-rich substrates and to minimize the accumulation of catabolic byproducts could partly explain the ability of the species to survive prolonged subzero conditions.

Haemolymph fluid osmolality influences the neutral-red retention assay in the eastern oyster

D. Méthé, L.A. Comeau, H. Stryhn, J.F. Burka, T. Landry and J. Davidson Department of Fisheries and Oceans, Gulf Fisheries Center, Science Branch, P.O. Box 5030, Moncton, New Brunswick, Canada E1C 9B6; Department of Health Management, Atlantic Veterinary College, University of Prince Edward Island, 550 University Avenue, Charlottetown, Prince Edward Island, Canada C1A 4P3; and Department of Biomedical Sciences, Atlantic Veterinary College, University of Prince Edward Island, 550 University Avenue, Charlottetown, Prince Edward Island, Canada C1A 4P3

DETERMINATION OF THE EFFECTS OF SEISMIC EXPLORATION ON FISH (PROJECT SEIFISH)

Of major international concern is the potential effect of marine seismic exploration on fish and fisheries. To date, only a few studies have directly examined this issue. In perhaps the most important set of studies, Engas et al. (1996) compared fish catch preand postseismic exposure and found a reduction in catch rates following seismic shooting. Slotte et al. (2004) did a similar study but watched fish movement with sonar and got results that suggested movement of fish to greater depths during seismic activity. However, to date, there are no detailed data on behaviour of individual fish in response to seismic activities. Such data are critical if investigators are to fully evaluate if and how seismic exploration impacts fisheries.