Lionel Camus

Akvaplan-niva, Tromsø, Troms, Norway

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Publications (60)153.14 Total impact

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    ABSTRACT: This study aimed to simulate conditions in which dispersant (Dasic NS) might be used to combat an oil spill in coastal sub-Arctic water of limited depth and water exchange in order to produce input data for Net Environmental Benefit Analysis (NEBA) of Arctic and sub-Arctic coastal areas. Concentration dependent differences in acute responses and long-term effects of a 48 h acute exposure to dispersed oil, with and without the application of a chemical dispersant, were assessed on the Arctic filter feeding bivalve Chlamys islandica. Icelandic scallops were exposed for 48 h to a range of spiked concentrations of mechanically and chemically dispersed oil. Short-term effects were assessed in terms of lysosomal membrane stability, superoxide dismutase, catalase, gluthatione S-transferases, glutathione peroxidases, glutathione reductase, glutathione, total oxyradical scavenging capacity, lipid peroxidation and peroxisomal proliferation. Post-exposure survival, growth and reproductive investment were followed for 2 months to evaluate any long-term consequence. Generally, similar effects were observed in scallops exposed to mechanically and chemically dispersed oil. Limited short-term effects were observed after 48 h, suggesting that a different timing would be required for measuring the possible onset of such effects. There was a concentration dependent increase in cumulative post-exposure mortality, but long-term effects on gonadosomatic index, somatic growth/condition factor did not differ among treatments.
    Full-text · Article · May 2016 · Ecotoxicology and Environmental Safety
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    ABSTRACT: In the context of an oil spill accident and the following oil spill response, much attention is given to the use of dispersants. Dispersants are used to disperse an oil slick from the sea surface into the water column generating a cloud of dispersed oil droplets. The main consequence is an increasing of the sea water-oil interface which induces an increase of the oil biodegradation. Hence, the use of dispersants can be effective in preventing oiling of sensitive coastal environments. Also, in case of an oil blowout from the seabed, subsea injection of dispersants may offer some benefits compared to containment and recovery of the oil or in situ burning operation at the sea surface. However, biological effects of dispersed oil are poorly understood for deep-sea species. Most effects studies on dispersed oil and also other oil-related compounds have been focusing on more shallow water species. This is the first approach to assess the sensitivity of a macro-benthic deep-sea organism to dispersed oil. This paper describes a toxicity test which was performed on the macro-benthic deep-sea amphipod (Eurythenes gryllus) to determine the concentration causing lethality to 50 % of test individuals (LC50) after an exposure to dispersed Brut Arabian Light (BAL) oil. The LC50 (24 h) was 101 and 24 mg L(-1) after 72 h and 12 mg L(-1) at 96 h. Based on EPA scale of toxicity categories to aquatic organisms, an LC50 (96 h) of 12 mg L(-1) indicates that the dispersed oil was slightly to moderately toxic to E. gryllus. As an attempt to compare our results to others, a literature study was performed. Due to limited amount of data available for dispersed oil and amphipods, information on other crustacean species and other oil-related compounds was also collected. Only one study on dispersed oil and amphipods was found, the LC50 value in this study was similar to the LC50 value of E. gryllus in our study. Since toxicity data are important input to risk assessment and net environmental benefit analyses, and since such data are generally lacking on deep-sea species, the data set produced in this study is of interest to the industry, stakeholders, environmental management, and ecotoxicologists. However, studies including more deep-sea species covering different functional groups are needed to evaluate the sensitivity of the deep-sea compartments to dispersed oil relative to other environmental compartments.
    No preview · Article · Dec 2015 · Environmental Science and Pollution Research
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    ABSTRACT: The decrease of ice cover in the Arctic will lead to an increase of ship traffic in the upcoming decades. Consequently, oil pollution is expected. In this context, the goals of this study were to evaluate the biological impact of marine diesel contamination and, on this basis, to determine analytical tools of interest (biomarkers) for future biomonitoring of diesel spills. Using a 7-day contamination protocol, this study investigated biochemical modulations in the digestive gland of the Iceland scallop (Chlamys islandica). Incorporation of contaminants was verified assessing haemolymph metabolites. Results showed a response of glutathione-S-transferase to contamination suggesting detoxification processes and the suitability of such a tool for diesel spill biomonitoring. The lack of modulation of superoxide dismutase activity and lipid peroxidation suggests no oxidative stress and the unsuitability of these molecular tools for biomonitoring.
    No preview · Article · Oct 2015 · Environmental Science and Pollution Research
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    Full-text · Dataset · Oct 2015
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    ABSTRACT: The current understanding of Arctic ecosystems is deeply rooted in the classical view of a bottom-up controlled system with strong physical forcing and seasonality in primary-production regimes. Consequently, the Arctic polar night is commonly disregarded as a time of year when biological activities are reduced to a minimum due to a reduced food supply. Here, based upon a multidisciplinary ecosystem-scale study from the polar night at 79°N, we present an entirely different view. Instead of an ecosystem that has entered a resting state, we document a system with high activity levels and biological interactions across most trophic levels. In some habitats, biological diversity and presence of juvenile stages were elevated in winter months compared to the more productive and sunlit periods. Ultimately, our results suggest a different perspective regarding ecosystem function that will be of importance for future environmental management and decision making, especially at a time when Arctic regions are experiencing accelerated environmental change [1].
    Full-text · Article · Oct 2015 · Current Biology
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    ABSTRACT: The influence of temperature on the bioaccumulation, toxicokinetics, biotransformation and depuration of pyrene was studied in the arctic marine amphipod Gammarus setosus. A two-compartment model was used to fit experimental values of total body burden, total metabolites and parent pyrene concentrations and to calculate toxicokinetic variables derived for two experimental treatments (2 and 8 °C). No statistically significant differences were observed with temperature for these toxicokinetic variables or bioconcentration factors. Contrarily, the Q10 values suggested that the toxicokinetic variables ke and km were temperature-dependent. This may be explained by the high standard deviation of the Q10 values. Q10 is the variation in the rate of a metabolic reaction with a 10 °C increase in temperature. Depuration rate constants were calculated from linear best fit equations applied to measured pyrene concentrations over time during the depuration phase of the experiment. During depuration, the parent pyrene was eliminated in two stages with faster elimination observed at 8 °C compared to 2 °C. This finding was also indicated by the Q10. No changes in total body burdens of metabolite concentrations were observed during the monitoring of depuration over a period of 96 h. The biotransformation pathway of pyrene in G. setosus was also investigated in this study with two main phase II biotransformation products discovered by liquid chromatography. These products are conditionally identified as the sulphate and glucose conjugates of 1-hydroxy-pyrene. Overall, the study contributes new knowledge to the understanding of the fate of PAHs in arctic biota. In particular, the study provides valuable insight into the bioaccumulation and biotransformation of an important PAH and its metabolites in a species that serves as both a predator and prey in the arctic ecosystem. Copyright © 2015 Elsevier Ltd. All rights reserved.
    No preview · Article · Aug 2015 · Marine environmental research
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    ABSTRACT: Concentration dependent differences in acute and long-term effects of a 48 h exposure to mechanically or chemically dispersed crude oil were assessed on juvenile lumpsucker (Cyclopterus lumpus). Acute or post-exposure mortality was only observed at oil concentrations representing higher concentrations than reported after real oil spills. Acute mortality was more apparent in chemically than mechanically dispersed oil treatments whereas comparable EC50s were observed for narcosis. There was a positive correlation between EROD activity and muscle PAH concentration for the lower oil concentrations whereas higher concentrations inhibited the enzyme activity. The incidence of gill tissue lesions was low with no difference between dispersion methods or oil concentrations. A concentration dependent decrease in swimming- and feeding behavior and in SGR was observed at the start of the post-exposure period, but with no differences between corresponding oil treatments. Three weeks post-exposure, fish from all treatments showed as high SGR as the control fish. Copyright © 2015 Elsevier Ltd. All rights reserved.
    Full-text · Article · Jan 2015 · Marine Environmental Research
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    ABSTRACT: Produced water is the main discharge stream from oil and gas production. For offshore activities this water is usually discharged to the marine environment. Produced water contains traces of hydrocarbons such as polycyclic aromatic hydrocarbons as well as alkylphenols, which are relatively resistant to biodegradation and have been reported to cause adverse effects to marine organisms in laboratory studies. For management of produced water, risk-based tools have been developed using toxicity data for mainly non-Arctic species. Reliable risk assessment approaches for Arctic environments are requested to manage potential impacts of produced water associated with increased oil and gas activities in Arctic regions. In order to assess the applicability of existing risk tools for Arctic areas, basic knowledge on the sensitivity of Arctic species has to be developed. In the present study, acute and chronic toxicity of artificial produced water for 6 Arctic and 6 temperate species was experimentally tested and evaluated. The hazardous concentrations affecting 5% and 50% of the species were calculated from species sensitivity distribution curves. Hazardous concentrations were compared to elucidate whether temperate toxicity data used in risk assessment are sufficiently representative for Arctic species. From the study it can be concluded that hazardous concentration derived from individual species' toxicity data of temperate and Arctic species are comparable. However, the manner in which Arctic and non-Arctic populations and communities respond to exposure levels above established thresholds remains to be investigated. Hence, responses at higher levels of biological organization should be studied to reveal potential differences in sensitivities to produced water between Arctic and non-Arctic ecosystems. Copyright © 2014 Elsevier Inc. All rights reserved.
    Full-text · Article · Dec 2014 · Ecotoxicology and Environmental Safety
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    ABSTRACT: Chemical dispersaiits are considered as an appropriate technical response to oil spills in offshore environments. Nevertheless, since dispersant can increase the bioavailability of oil. its use is still a controversial issue. Furthermore, there is a lack of information related to the ecotoxicity of the oil-dispersant mixture especially for the Arctic area. Therefore, the aim of this study was to compare the impact on tissue respiration of a dispersed oil (weathered Crude Arabian Light) 011 two fish species, sea bass (.Dicentrarchus labrax) and polar cod (Boreogadus saida) representative respectively of temperate and Arctic water ecosystems. Polar cod (n= 32: 94 ± 4 g) and sea bass (n= 32: 492 ± 20 g) were exposed for 48 hours to one of the following treatments: control, mechanically dispersed oil. chemically dispersed oil and dispersant alone. The impacts of these exposure conditions were assessed 011 heart energy metabolism using respirometry on permeabilized cardiac fibers. Following exposure, alteration in measurements of O2 consumption by permeabilized cardiac fibers was found for the two species. The results show that for polar cod. oil alone decreased the activity of the respiratory chain (0.35 ± 0.05 02.min-1.mg of dry tissues-1 for control group: 0.20 ± 0.03 CKO2min-1mg of dry tissues-1 for mechanically dispersed oil group) whereas the dispersant alone did not have any impact (0.30 ± 0.05 O2.min-1.mg of dry tissues-1 for dispersant alone group). For sea bass, the results were different, dispersant alone decreased the activity of the respiratory chain (1.15 ± 0.11 O2min-1mg of dry tissues-1 for control group: 0.47 ± 0.04 O2min-1mg.mg of dry tissues-1 for dispersant alone group) whereas the results for oil alone were not different from the control group (1.26 ± 0.32 O2min-1mg of dry tissues-1 for mechanically dispersed oil group). These results show that oil and dispersaiits can alter mitochondrial activity and are a clear demonstration of the inter-specific variation in risk assessment.
    No preview · Conference Paper · Jun 2014
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    ABSTRACT: Lysosomal autophagic responses, such as lysosomal membrane stability, neutral lipids (NL), lipofuscin (LF), and malondialdehyde (MDA) levels, are valuable measures of cellular early-onset effects induced by environmental stress factors, such as contaminant exposure and fasting. In this study, these parameters were analysed and related to levels of halogenated organic contaminants (HOCs) in 40 Herring Gull (Larus argentatus) chicks. Chicks were experimentally exposed to HOCs through diet and went through a period of nutrient deprivation at the end of the experiment. HOC exposure and fasting were conducted separately and in combination. NL storages were depleted, and lysosomal membranes were destabilised after HOC exposure and nutrient deprivation. These responses were not related specifically to one type of stress or the extent of the treatment. No synergistic or additive effects from the combination of HOC exposure and fasting were observed. LF accumulated, and MDA levels increased as a result of fasting, but were unaffected by HOC exposure. LF accumulation was strongly associated with the percent weight change in the chicks. Large weight loss was associated with high LF levels, and slight weight gain was associated with low LF levels. Hence, food deprivation affected all the measured parameters, and HOC exposure decreased NL levels and lysosomal membrane stability in HG chick liver. Furthermore, autophagic lysosomal parameters have frequently been applied as biomarkers of cellular health status in previous studies of marine and terrestrial invertebrates, and this study suggests that these parameters may be good candidates for biomarkers of cellular health status in seabirds as well.
    Full-text · Article · Apr 2014 · Comparative Biochemistry and Physiology Part C Toxicology & Pharmacology
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    ABSTRACT: Offshore oil and gas drilling processes generate operational discharges such as produced water (PW), a complex mixture of seawater with polycyclic aromatic hydrocarbons (PAH) and alkylphenols (AP). Some of these compounds may interact with the endocrine system of marine organisms and alter reproductive functions. In this study, polar cod were exposed for up to 28 d to a mixture of PAH, alkylated PAH, and AP simulating the composition of North Sea PW, at low and high concentrations (1:2000 and 1:1000 dilution of the original concentrate, respectively). Potential adverse effects of PW on polar cod physiology were investigated through biomarkers of biotransformation (hepatic ethoxyresorufin O-deethylase [EROD] activity and bile PAH metabolites), endocrine disruption (plasma vitellogenin [VTG] levels and sex steroid concentrations), and gonad histology. Plasma sexual steroid levels in fish were not markedly affected by PW exposure, while higher plasma VTG concentrations were measured in females exposed to the high PW treatment for 7 and 28 d. In males exposed to the higher PW concentration, inhibition of spermatogenesis was observed after 28 d in addition to increase of melano-macrophage occurrence in testis. Females exposed to the high PW treatment for 21 d showed a significant increase of atresia incidence. Finally, a significant decrease in oocyte number was observed in high PW exposed female ovaries after 28 d of exposure.
    Full-text · Article · Apr 2014 · Journal of Toxicology and Environmental Health Part A
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    ABSTRACT: In this study, impact of dispersed oil on cardiac mitochondrial function was assessed in a key species of Arctic marine ecosystem, the polar cod Boreogadus saida. Mature polar cod were exposed during 48 h to dispersed oil (mechanically and chemically) and dispersants alone. The increase observed in ethoxyresorufin-O-deethylase activity and polycyclic aromatic hydrocarbon metabolites in bile indicated no difference in contamination level between fish exposed to chemical or mechanical dispersion of oil. Oil induced alterations of O2 consumption of permeabilised cardiac fibres showing inhibitions of complexes I and IV of the respiratory chain. Oil did not induce any modification of mitochondrial proton leak. Dispersants did not induce alteration of mitochondrial activity and did not increase oil toxicity. These data suggest that oil exposure may limit the fitness of polar cod and consequently could lead to major disruption in the energy flow of polar ecosystem.
    No preview · Article · Feb 2014 · Environmental Science and Pollution Research
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    Full-text · Article · Jan 2014
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    ABSTRACT: Biologically treated wastewater (WW) from the Hammerfest LNG (liquefied natural gas) plant is discharged to the sea. A study using biomarkers in mussels and Atlantic cod was performed to examine whether this discharge meets a zero harmful emission requirement. Caging of mussels close to the outfall and exposure of mussels and fish to WW in the laboratory were conducted, and a suite of contaminant responsive markers was assessed in exposed animals. In mussels the markers included chemical contaminant levels, haemocyte lysosomal instability and nucleus integrity, cellular energy allocation, digestive gland and gonad histopathology and shell-opening behaviour. In fish, biliary PAH metabolites and gill histopathology biomarkers were measured. A consistent cause-effect relationship between WW treatments and markers measured in test animals was not found. The results therefore indicate that the WW emission is unlikely to represent a significant stress factor for the local marine environment under the conditions studied.
    Full-text · Article · Feb 2013 · Marine Pollution Bulletin
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    ABSTRACT: Due to a northward shift in oil and gas activities, there is an increasing need to understand the potential anthropogenic impacts of oil-related compounds on sub-Arctic and Arctic organisms, particularly those in coastal habitats. Capelin (Mallotus villosus), a key fish species in the Barents Sea ecosystem, undertakes aggregated spawning at both intertidal and subtidal coastal localities in northern Norway. To investigate the sensitivity of capelin embryos to oil compounds, newly fertilized capelin eggs were collected from a spawning beach and exposed until hatch (32 days) to either the water soluble fraction of crude oil or the single PAH compound, pyrene. Threshold levels for egg mortality, development and hatching success were determined. Concentrations of 40 μg/L crude oil (∑26 PAHs) and 55 μg/L pyrene significantly increased embryonic mortality rates and decreased hatching success, compared with controls, indicating that a potential oil spill in the vicinity of capelin spawning grounds may cause significant impacts. No significant incidence of adverse effects such as yolk sac oedema, pericardia oedema, haemorrhages, craniofacial abnormalities, premature hatch or inhibited growth was observed. Histological studies of hatched larvae did not reveal specific sublethal effects in tissues and organs. Developmental delays and subsequent embryo death were noticed at the period of eye pigmentation in affected groups. Early life-history stages of capelin are sensitive indicators of PAH impacts, but the mechanisms responsible for the toxic effects require further investigation.
    Full-text · Article · Feb 2012 · Aquatic toxicology (Amsterdam, Netherlands)
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    ABSTRACT: In the Barents Sea, the limited data on biological relevant indicators and their responses to various anthropogenic stressors have hindered the development of a consistent scientific basis for selecting indicator species and developing practical procedures for environmental monitoring. Accordingly, the main aim of the present study was to develop a common set of baseline values for contaminants and biomarkers in three species, and to identify their strengths and limitations in monitoring of the Barents Sea. Blue mussel (Mytilus edulis), Icelandic scallop (Chlamys islandica) and Atlantic cod (Gadus morhua) were sampled from a north Norwegian fjord in March, June, September and December 2010. Digestive glands from the bivalve species and liver from Atlantic cod were analysed for biomarkers of oxidative stress (catalase [CAT], glutathione peroxidase [GPX], glutathione-S-transferase activities [GST], lipid peroxidation as thiobarbituric reactive substances [TBARS] and total oxyradical scavenging capacity [TOSC]), biotransformation (ethoxyresorufine-O-deethylase activity [EROD]) and general stress (lysosomal membrane stability [LMS]). Concentrations of polycyclic aromatic hydrocarbons (PAHs) and metals in the bivalves and PAH metabolites in fish bile were quantified. Finally, energy reserves (total lipids, proteins and carbohydrates) and electron transport system (ETS) activity in the digestive gland of the bivalves and liver of Atlantic cod provided background information for reproductive cycle and general physiological status of the organisms. Blue mussel and Icelandic scallop showed very similar trends in biological cycle, biomarker expression and seasonality. Biomarker baselines in Atlantic cod showed weaker seasonal variability. However, important biological events may have been undetected due to the large time intervals between sampling occasions. Physiological biomarkers such as energy reserves and ETS activity were recommended as complementary parameters to the commonly used stress biomarkers, as they provided valuable information on the physiological status of the studied organisms. Interpretation of the seasonality in oxidative stress biomarkers was in general difficult but TOSC and lipid peroxidation were preferred over the antioxidant enzyme activities. This study is the first reporting seasonal baseline in these three species in a sub-Arctic location. Overall, the Icelandic scallop was considered the most adequate organism for environmental monitoring in the Barents Sea due to the interpretability of the biomarker data as well as its abundance, ease to handle and wide distribution from the southern Barents Sea to Svalbard.
    Full-text · Article · Jan 2012 · Aquatic toxicology (Amsterdam, Netherlands)
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    ABSTRACT: Reliable risk assessment approaches for Arctic environments are requested to manage potential impacts associated with increased activities in Arctic regions. We performed toxicity tests on Arctic and temperate species exposed to the narcotic acting oil component, 2-methyl naphthalene. The experimental results were used to quantify concentration causing lethality to 50% of exposed individuals and no-effect concentration (individual level). For estimates at community level, the hazardous concentrations affecting 5% and 50% of the species were calculated from sensitivity distribution curves. These survival metrics were then used to elucidate whether temperate toxicity data used in risk assessment are sufficiently representative for the Arctic. Taking data uncertainty into consideration, we found no regional difference in tolerances to 2-methyl naphthalene either at the species level or at the community level. Hence these data support a conclusion that values of survival metrics for temperate regions are transferrable to the Arctic for the chemical 2-methyl naphthalene, as long as extrapolation techniques are properly applied and uncertainties are taken into consideration.
    Full-text · Article · Aug 2011 · Marine environmental research
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    ABSTRACT: Lysosomal membrane stability, lipofuscin (LF), malondialdehyde (MDA), neutral lipid (NL) levels, as well as halogenated organic compounds (HOCs), Cr, Cd, Pb and Fe concentrations were analyzed in liver of black-legged kittiwake (BK), herring gull (HG), and northern fulmar (NF) chicks. There were significant species differences in the levels of NL, LF and lysosomal membrane stability. These parameters were not associated with the respective HOC concentrations. LF accumulation was associated with increasing Cr, Cd and Pb concentrations. HG presented the lowest lysosomal membrane stability and the highest. LF and NL levels, which indicated impaired lysosomes in HG compared to NF and BK. Lipid peroxidation was associated with HOC and Fe2+ levels. Specific HOCs showed positive and significant correlations with MDA levels in HG. The study indicates that contaminant exposure can affect lysosomal and lipid associated parameters in seabird chicks even at low exposure levels. These parameters may be suitable markers of contaminant induced stress in arctic seabirds.
    Full-text · Article · Jul 2011 · Marine Pollution Bulletin
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    ABSTRACT: The Herring gull (HG) (Larus argentatus) is naturally exposed to halogenated organic compounds such as polychlorinated biphenyls (PCBs) through its diet. During periods of food scarcity, arctic seabirds experience lipid mobilization, allowing stored lipid soluble contaminants to re-enter the body circulation. In the present study, we investigated the effects of PCB exposure and fasting on the antioxidant defense system in HG chicks. Forty newly hatched chicks were exposed to contaminated cod liver oil for 6weeks and then fasted for 1week. We assessed the hepatic total oxyradical scavenging capacity (TOSC) against peroxynitrite, hydroxyl and peroxyl radicals, and measured glutathione (reduced: GSH, and oxidized: GSSG) levels and the enzymatic activities of catalase, glutathione peroxidase, glutathione reductase and superoxide dismutase. The results show that fasting significantly increased the HOC levels in the HG chick livers. Limited effects were observed on antioxidant responses; significant effects were only found for catalase (CAT) activity, Se-dependent GPX activity and the GSH/GSSG ratio in the exposed and fasted group. CAT and Se-dependent GPX activities correlated negatively with the PCB concentrations within this group, and a nonlinear relationship between glutathione and contaminant levels was also found. These effects were generally not observed after exposure or fasting alone and were likely related to the high PCB levels induced by the combination of exposure and fasting.
    Full-text · Article · Jun 2011 · Science of The Total Environment
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    ABSTRACT: The past decades of monitoring discharges from oil and gas industry have revealed that although there are indications of adverse effects in tissues of aquatic organisms, little is known about their temporal development. Furthermore, observations in wild-caught individuals have not been clearly reproduced in laboratory studies or caging studies, and vice versa, and the results are therefore not easily interpretable. There is clearly a need for exposure studies designed for monitoring the development of effect markers in individual fish over chronic periods to low contaminant levels. Through repetitive nondestructive sampling, the progression of effects may be monitored in individuals, significantly reducing the number of fish needed in exposure studies. A laboratory exposure study was designed to be able to monitor selected parameters in individual Atlantic cod (Gadus morhua). Passive integrated transponders in combination with visible implant elastomers were used to study individual fish during the exposure period (44 wk). Fish were measured (weight and length) and a blood sample was taken for analysis of hematocrit, DNA damage (micronucleus), and oxidative stress (total oxyradical scavenging capacity) at up to seven time points. There were no apparent adverse effects of treatments on the health of experimental fish, frequency of micronucleated erythrocytes, or oxidative stress in whole blood. It is possible that the time scale was not sufficient for development and detection of parameters included here or that red blood cells may not be a suitable matrix for the selected analyses. Future studies need to include other parameters in blood to investigate their sensitivity to low-concentration exposures.
    Full-text · Article · Mar 2011 · Journal of Toxicology and Environmental Health Part A

Publication Stats

871 Citations
153.14 Total Impact Points

Institutions

  • 2007-2015
    • Akvaplan-niva
      Tromsø, Troms, Norway
  • 2004-2015
    • University Centre in Svalbard (UNIS)
      Longyearbyen, Svalbard, Svalbard and Jan Mayen
  • 2008
    • International Research Institute of Stavanger
      Stavenger, Rogaland, Norway
  • 2002-2004
    • University of Plymouth
      • School of Biological Sciences
      Plymouth, England, United Kingdom