CO2 budget of cultured mussels metabolism in the highly productive Northwest Iberian upwelling system.

Assessing the carbon footprint of marine bivalve aquaculture demands an accurate estimation of the CO2 release associated to capital goods and aquaculture operations but also to the metabolic CO2 budget of the cultured species. Nowadays, there are discrepancies on the processes to include in that budget, how to estimate them, and which scale should be applied, from individual to ecosystem. Site-specific environmental conditions and culture methods also affect significantly the estimates. Here, we have gathered environmental, biochemical and metabolic data from published scientific articles, reports and existing databases to present the metabolic CO2 budget for mussel aquaculture in the coastal inlets of the Northwest Iberian upwelling. We analyse the contribution of mussel flesh and shell production jointly and separately. At the individual scale, the shell CO2 budget is estimated from CO2 removal by shell matrix protein synthesis and CO2 release during calcification and respiration to support shell maintenance. Organic carbon in mussel flesh and CO2 released by respiration to support flesh maintenance contribute to the flesh CO2 budget. Only calcification and respiration processes are considered when estimating the metabolic carbon footprint of individual mussels because organic carbon in mussel flesh and shell returns to the atmosphere as CO2 in a relatively short period. While the metabolic carbon footprint associated to mussel shell remains constant at 365 kg CO2 per ton of shell, it varies from 92 to 578 kg CO2 per ton of mussel flesh. This large variability depends on mussel seeding time and harvesting size, due to the differential seasonal growth patterns of flesh and shell. Inclusion of the CO2 potentially immobilised in mussel faeces buried in the sediments would lead to a reduction of the metabolic carbon footprint estimates by up to 6 % compared with the individual estimates.

Modeling the impact of climate change on mussel aquaculture in a coastal upwelling system: A critical assessment.

Forecasting of climate change impacts on marine aquaculture production has become a major research task, which requires taking into account the biases and uncertainties arising from ocean climate models in coastal areas, as well as considering culture management strategies. Focusing on the suspended mussel culture in the NW Iberian coastal upwelling system, we simulated current and future mussel growth by means of a multistructural net production Dynamic Energy Budget (DEB) model. We considered two scenarios and three ocean climate models to account for climate uncertainty, and applied a bias correction to the climate models in coastal areas. Our results show that the predicted impact of climate change on mussel growth is low compared with the role of the seeding time. However, the response of mussels varied across climate models, ranging from a minor growth decline to a moderate growth increase. Therefore, this work confirms that an accurate forecasting of climate change impacts on shellfish aquaculture should take into account the variability linked to both management strategies and climate uncertainty.

Characterizing individual variability in mussel (Mytilus galloprovincialis) growth and testing its physiological drivers using Functional Data Analysis.

Determining the magnitude and causes of intrinsic variability is a main issue in the analysis of bivalve growth. Inter-individual variability in bivalve growth has been attributed to differences in the physiological performance. This hypothesis has been commonly tested comparing the physiological rates of fast and slow growers after size differentiation has occurred. This experimental design may detect a link between growth and physiological performance, but we cannot interpret the posterior physiological performance as a driver for the prior growth variability. Considering these limitations, this work introduces a new methodological framework for the analysis of bivalve growth variability. We have conducted sequential measurements of size and physiological performance (feeding, digestion and metabolic rates) in even-sized mussels growing under homogeneous environmental conditions. This experimental design allows us to distinguish between changes over time within individuals, i.e. growth and trends in the physiological rates, from differences between individuals with respect to a baseline level. In addition, Functional Data Analysis provides powerful tools to summarize all the information obtained in the exhaustive sampling scheme and to test whether differences in the physiological performance enhance growth dispersion. Our results report an increasing dispersion in both size and physiological performance over time. Although mussels grew during the experiment, it is difficult to detect any increasing or decreasing temporal pattern in their feeding, digestion and metabolic rates due to the large inter-individual variability. Comparison between the growth and physiological patterns of mussels with final size above (fast growers) and below (slow growers) the median found that fast growers had larger feeding and digestion rates and lower metabolic expenditures during the experimental culture than mussels with slow growth, which agrees with the hypothesis of a physiological basis for bivalve growth variability.

From classical to nonparametric growth models: Towards comprehensive modelling of mussel growth patterns.

Understanding biological processes, such as growth, is crucial to development management and sustainability plans for bivalve populations. Von Bertalanffy and Gompertz models have been commonly used to fit bivalve growth. These models assume that individual growth is only determined by size, overlooking the effects of environmental and intrinsic conditions on growth patterns. The comparison between classical models and nonparametric GAM (generalized additive models) fits conducted in this work shows that the latter provide a more realistic approach of mussel growth measured in terms of shell length, and dry weight of hard and soft tissues. GAM fits detected a reduction in growth during the cold season, under unfavourable nutritional conditions. These fits also captured the decoupling between hard and soft tissue growth, widely addressed in the literature but not incorporated in growth models. In addition a GAM fit of condition index allowed us to explain annual changes in resources allocation, identifying the asymptotic growth of shell and the effects of the reproductive cycle on soft tissue fluctuations.

Solar irradiance dictates settlement timing and intensity of marine mussels.

Identifying the environmental factors driving larval settlement processes is crucial to understand the population dynamics of marine invertebrates. This work aims to go a step ahead and predict larval presence and intensity. For this purpose we consider the influence of solar irradiance, wind regime and continental runoff on the settlement processes. For the first time, we conducted a 5-years weekly monitoring of Mytilus galloprovincialis settlement on artificial suspended substrates, which allowed us to search for interannual variability in the settlement patterns. Comparison between the seasonal pattern of larval settlement and solar irradiance, as well as the well-known effect of solar irradiance on water temperature and food availability, suggest that solar irradiance indirectly influences the settlement process, and support the use of this meteorological variable to predict settlement occurrence. Our results show that solar irradiance allows predicting the beginning and end of the settlement cycle a month in advance: Particularly we have observed that solar irradiance during late winter indirectly drives the timing and intensity of the settlement onset, Finally, a functional generalise additive model, which considers the influence of solar irradiance and continental runoff on the settlement process, provides an accurate prediction of settlement intensity a fortnight in advance.

Flexibility of Physiological Traits Underlying Inter-Individual Growth Differences in Intertidal and Subtidal Mussels Mytilusgalloprovincialis.

Mussel seed (Mytilusgalloprovincialis) gathered from the intertidal and subtidal environments of a Galician embayment (NW, Spain) were maintained in the laboratory during five months to select fast (F) and slow (S) growing mussels. The physiological basis underlying inter-individual growth variations were compared for F and S mussels from both origins. Fast growing seemed to be a consequence of greater energy intake (20% higher clearance and ingestion rate) and higher food absorption rate coupled with low metabolic costs. The enhanced energy absorption (around 65% higher) resulted in 3 times higher Scope for Growth in F mussels (20.5±4.9 J h(-1)) than S individuals (7.3±1.1 J h(-1)). The higher clearance rate of F mussels appears to be linked with larger gill filtration surface compared to S mussels. Intertidal mussels showed higher food acquisition and absorption per mg of organic weight (i.e. mass-specific standardization) than subtidal mussels under the optimal feeding conditions of the laboratory. However, the enhanced feeding and digestive rates were not enough to compensate for the initial differences in tissue weight between mussels of similar shell length collected from the intertidal and subtidal environments. At the end of the experiment, subtidal individuals had higher gill efficiency, which probably lead to higher total feeding and absorption rates relative to intertidal individuals.

Contrasting physiological responses of two populations of the razor clam Tagelus dombeii with different histories of exposure to paralytic shellfish poisoning (PSP).

This study describes the physiological performance of two populations of the razor clam Tagelus dombeii from two geographic areas with different histories of exposure to paralytic shellfish poisoning (PSP) linked to the toxic dinoflagellate Alexandrium catenella. Clams from Melinka-Aysén, which are frequently exposed to PSP, were not affected by the presence of toxins in the diet. However, clams from Corral-Valdivia, which have never been exposed to PSP, exhibited significantly reduced filtration activity and absorption, affecting the energy allocated to scope for growth (SFG). Ammonia excretion and oxygen uptake were not affected significantly by the presence of A. catenella in the diet. Measurements of energy acquisition and expenditure were performed during a 12-day intoxication period. According to three-way repeated measure ANOVAs, the origin of the clams had a highly significant effect on all physiological variables, and the interaction between diet and origin was significant for the clearance and absorption rates and for the scope for growth. The scope for growth index showed similar positive values for both the toxic and non-toxic individuals from the Melinka-Aysén population. However, it was significantly reduced in individuals from Corral-Valdivia when exposed to the diet containing A. catenella. The absence of differences between the physiological response of the toxic and non-toxic clams from Melinka-Aysén may be related to the frequent presence of A. catenella in the environment, indicating that this bivalve does not suffer negative consequences from PSP. By contrast, A. catenella has a negative effect on the physiological performance, primarily on the energy gained from the environment, on T. dombeii from Corral-Valdivia. This study supports the hypothesis that the history of PSP exposure plays an important role in the physiological performance and fitness of filter feeding bivalves.

Fatty acids as tracers of trophic interactions between seston, mussels and biodeposits in a coastal embayment of mussel rafts in the proximity of fish cages.

We traced the food sources of mussel Mytilus galloprovincialis cultured in suspension in Ría Ares-Betanzos (N.W. Spain) by means of fatty acid (FA) biomarkers. The FA profile of seston, mussels' mantle, digestive gland and feces was analyzed during five seasons. Due to the proximity of a fish farm to the bivalve aquaculture site, we also tested if mussels and seston situated 170 m distant from the fish cages incorporated fish feed FA markers compared with samples obtained 550 m away. The principal FA in the mussels' organs were 16:0, 16:1ω7, EPA (20:5ω3) and DHA (22:6ω3), while 16:0 predominated in the feces. Seasonal fluctuations in the seston composition were mirrored in the FA signature of mussels' organs and feces, although the digestive gland had the closest resemblance to the seston FA profile. In general, diatom and bacteria derived-biomarkers predominated in mussels' organs and feces during the upwelling period (spring-summer), while dinoflagellates were the dominant dietary source during downwelling (autumn-winter). The higher concentration of EPA and DHA in both organs and the feces compared with the seston suggested a preferential accumulation of these ω3 FA in the mussels' tissues. The results showed a lack of assimilation of fish feed FA biomarkers in the seston and mussel samples. This might be due to the dispersion of uneaten feed particles by high current velocity, substantial distance between the fish and mussel culture, the limited amount of nutrient waste released by the fish farm and dilution of feed particles in the large mussel standing stock.

Variability in biochemical components of the mussel (Mytilus galloprovincialis) cultured after Prestige oil spill.

The biochemical composition (proteins, carbohydrates, glycogen, total lipids and lipid classes) of the mussel Mytilus galloprovincialis was investigated during an experimental culture using mussel seed from areas with different degree of exposure to the Prestige oil spill. The aim of the study was to identify alterations in the biochemical composition of mussel seed from natural populations commonly used in Galicia for mussel raft culture that might be linked to previous oil exposure. We have selected three mussel seed populations from Pindo, Miranda and Redes, that were characterised in a previous study according to the oil exposure three months after the spill. These populations were transplanted to a raft culture system in the Ría de Ares-Betanzos where our experimental culture followed standard commercial techniques from March 2003 to February 2004. Mussels from Pindo (characterised as the most affected area by the oil spill) showed marked differences in lipid composition with regard to other populations in the content of triacylglycerols, (P<0.001), free fatty acids (P<0.001) and phospholipids (P<0.05) at the onset of the culture. Although these differences in lipid composition might reflect their previous exposition to hydrocarbons, this pattern did not last longer most likely due to depuration of hydrocarbons stored in the tissues or by the development of certain tolerance to PAHs. These significant differences were not detected between Miranda (designed as hardly affected area) and Redes (designed as reference area) which may reflect that Miranda mussels were not affected or only hardly affected by the spill. With the exception of the onset of the culture, biochemical composition showed similar patterns in all mussel populations. Then, the fact of being cultured in a common environment seemed to be more responsible for the long-term variability in the energetic reserve than the origin of the populations or their previous biochemical status.

Free amino acid composition in juveniles of Mytilus galloprovincialis: spatial variability after Prestige oil spill.

Composition of free amino acids (FAA) in juveniles of Mytilus galloprovincialis was analysed along a large geographical coastline area in Galicia (NW Spain). Individuals were sampled in February 2003, three months after the Prestige oil spill. Pollution values at sampling time were reported as polycyclic aromatic hydrocarbons (PAHs) concentrations in soft tissues of individuals and varied between the highest amount observed in Carrumeiro mussels (502 ng/g dw) and the lowest in Pindo mussels (196 ng/g dw), both locations being close to each other in the centre of the geographical area under study. Pollution values in the other populations varied within the range of 241-347 ng/g dw. Total free amino acids (TFAA) were highest in Aguiño-Pindo-Carrumeiro juveniles at the centre of the Coastline area studied (420-462 micromol/g dw) as compared to the other populations at North and South of Galicia (312-347 micromol/g dw). TFAA results were based on the variability observed in protein free amino acids (PFAA micromol/g dw) among populations (214-249 micromol/g dw for Aguiño-Pindo-Carrumeiro mussels and 98-149 micromol/g dw for the other populations) whereas non-protein free amino acids (NPFAA) taurine and ornithine did not show any significant spatial pattern of variation. Glycine and alanine represented the most abundant PFAA (16-29% and 2.7-11.9% of TPFAA, respectively) and significant correlations between PFAA and both the protein content of soft tissues (r=-0.82) and the condition index of juveniles (r=0.86) were observed. No significant relationships were detected, however, between pollution values in soft tissues as PAHs and FAA profiles with the exception of alanine concentrations as percentage of TFAA (r=0.88; P<0.01). The latter seemed to be an "all or nothing" effect likely due to the influence of other abiotic factors at one of the sampling sites. Such relationship was found not significant when the outlier represented by Carrumeiro mussels was removed from the analysis. The most abundant free amino acid taurine (43.2-68.5%TFAA) followed an inverse variability of that of glycine and by extension of the group PFAA most likely as a compensatory decrease in mussel populations with low protein content (and high condition index). Accordingly, taurine:glycine (t:g) ratio varied between 1 and 2 in most mussel populations but increased up to 3.2-4.2 in Miranda and Bueu mussels at both ends of the geographical interval studied with a corresponding PAHs concentrations of 261 and 304 ng/g dw, respectively. These mussel populations with the highest t:g ratios were characterised by the lowest PFAA contents (below 40%) and condition index values (below 10%). Results of the present study established a significant link between energetic status of growing juveniles and FAA concentrations in environments with different pollution degrees. Variability of the free amino acids profiles in soft tissues were related to endogenous factors of juveniles (protein content, condition index) whereas no relationship with contamination values could be observed. The utility of t:g ratio as general condition factor for M. galloprovincialis is also corroborated for in situ growing juveniles.