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The distributions of phytoplankton, zooplankton and hydrographic features off the coast of northern Norway were assessed in late April – early May 2019 using ship-based observations (CTD casts and Moving Vessel Profilers) and autonomous vehicles. A satellite chlorophyll climatology was generated to place our in-situ observations within a longer temporal sequence. Substantial spatial and temporal variability on all scales was observed in both the observations and climatology. Spring phytoplankton accumulation usually is initiated in the south on the continental shelf, and advanced in a northerly direction through time. Accumulations in the surface layer of deeper waters off the continental shelf occurred 2–3 weeks later than those on the shelf. During our survey, primary productivity was greatest in offshore waters where nutrients were not depleted and exceeded 2 g C m−2 d−1. The greatest Calanus finmarchicus abundances were associated with low chlorophyll concentrations, suggesting a major impact of grazing on phytoplankton biomass, but estimates of phytoplankton growth and zooplankton removal suggested that Calanus was responsible for a variable fraction (3–69%) of the daily chlorophyll changes. Vertical changes in chlorophyll were related to physical features during some transects, but to grazing and sinking in others. Understanding the spatial and temporal variations of the coupling of phytoplankton to zooplankton is essential to effective management of this important commercial species in Norwegian waters. • Phytoplankton primary productivity was assessed in northern Norway continental shelf and slope. • Biomass and productivity were greatest over the continental slope. • A satellite climatology showed that blooms spread from continental shelf to deep water. • Phytoplankton vertical distributions were controlled by multiple processes. • Copepods and chlorophyll were largely inversely correlated. [ABSTRACT FROM AUTHOR]

• Selective copepod grazing and water mass origin impacted spring bloom composition. • Diatom bloom enhanced zooplankton recruitment and deep carbon export. • Spring bloom composition impacted summer plankton community. • Mixo- and heterotrophic protists dominated the nutrient-poor summer months. • Copepod grazers controlled the summer protist community. Seasonal plankton time-series data are presented from Kongsfjorden from two years with contrasting environmental conditions. Kongsfjorden (west coast of Spitsbergen – 79°N) integrates inputs from Atlantic and Arctic waters, and glacier run-off, and is thus a prime location to study impacts on ecosystem dynamics of key environmental drivers that are relevant across the Arctic. Despite extensive research in Kongsfjorden, seasonally-resolved data are scarce. From late April/early May to early September 2019 and 2020, we conducted pelagic sampling at a mid-fjord station at mostly weekly to bi-weekly resolution investigating the environmental drivers of phyto- and zooplankton community composition and phenology. During spring 2019, Atlantic water masses with temperatures > 1 °C were found throughout the upper 250 m of the water column, and little sea ice occurred in the fjord. Spring 2020, in turn, was characterized by the presence of local water masses with sub-zero temperatures and relatively extensive sea-ice cover. The most striking contrast between the two years was the difference in phytoplankton spring bloom composition. In 2019, the spring bloom was dominated by the colonial stage of the haptophyte Phaeocystis pouchetii and diatoms played a minor role , while the spring bloom in 2020 was dominated by diatoms of the genus Thalassiosira succeeded by P. pouchetii. Selective grazing by large copepods and water mass structure seem to have been the decisive factors explaining the marked difference in diatom spring bloom biomass between the years while similar spring abundances of P. pouchetii in both years indicated that this species was less impacted by those factors. Our data suggest that differences in spring bloom composition impacted trophic transfer and carbon export. Recruitment of the dominant copepods Calanus finmarchicus and C. glacialis , Cirripedia and euphausiid larvae as well as the export of carbon to the seabed was more efficient during the diatom-dominated compared to the P. pouchetii– dominated spring bloom. In summer, the plankton composition shifted towards a flagellate-dominated community characterized by mixo- and heterotrophic taxa adapted to a lower nutrient regime and strong top-down control by copepod grazers. However, residual silicic acid after the P. pouchetii– dominated spring bloom fueled a late summer diatom bloom in 2019. Our data provide a first glimpse into the environmental drivers of plankton phenology and underline that high-resolution monitoring over many annual cycles is required to resolve the ephemeral variations of plankton populations against the backdrop of climate change. [ABSTRACT FROM AUTHOR]

The growth of Calanus finmarchicus nauplii was studied through three spawning seasons (2007, 2009 and 2011) using flow-through tubes. Natural seston was collected every second day and added as food source using a peristaltic pump. A mixture of cultured microalgae supplied in surplus concentration was used as a control treatment. No significant correlation between growth and food concentration measured as Chlorophyll a (Chla) or particulate organic carbon (POC) was detected, but the growth rate was significantly related to the content of EPA (20:5n-3) and DHA (22:6n-3) in the seston. The growth rate was overall higher for nauplii fed cultured microalgae compared to the nauplii fed natural seston. Although the nauplii fed algae cultures were fed surplus food, the growth did vary between the growth periods. Furthermore, the growth rate for nauplii fed natural seston and for nauplii fed cultured algae were positively related, suggesting that the maternal condition and the food quality experienced by the mothers could explain some of the variation in naupliar growth rate. We present lipid class data on Calanus finmarchicus eggs from field samples that, contrary to previous studies, showed a high content of wax esters. Fatty acid analyzes of eggs, nauplii stages and copepodites from field samples showed that eggs and nauplii have a similar fatty acid composition and that the main increase in the content and share of DHA and EPA was from nauplii to copepodite. The secondary production measured as naupliar growth was compared to the secondary production measured as carbon specific female egg production rate. The secondary production measured as egg production was generally higher than the secondary production measured as naupliar growth early in the spring, whereas the opposite situation was observed during post-bloom situations in late spring/early summer.

The growing world population requires large, renewable sources of nutritional food. Fish and other marine resources are nutrient dense and rich in healthy marine omega-3 lipids (EPA and DHA) beneficial to humans and animals alike. However, only about 6% of current fisheries are underexploited. Although improvements in fisheries management can increase marine production, aquaculture and untapped marine resources may contribute to a greater degree. Mesopelagic fish have increasingly gained interest as a massive unexploited marine resource. We did a literature review considering the narratives of mesopelagics (especially fish) as a new source of marine resource and compared this to similar narratives about the zooplankton redfeed (Calanus finmarchicus) - another marine resource that was considered equal in terms of potential. We found that mesopelagics have the potential to contribute to food production through usage in fish feed, but more significantly by direct human consumption. However, proper markets and demand must be present for them to make harvest economically viable. In addition, a thorough knowledge base will need to be generated to understand all the risks and make the harvest sustainable, in order to avoid adverse effects on several important species relying on mesopelagics as food. As technological advancements and ecological knowledge are increasing, with a growing focus on sustainable resources and healthy oceans as per SDG 14 and the UN Ocean Decade, time will tell whether your future diet will consist of mesopelagic fish, granted we are able to find and sustainably harvest the elusive treasure hidden in the twilight depths. • New marine resources can increase the output from the world's ocean. • Mesopelagics are the world's largest unexploited resource. • The zooplankton redfeed was previously considered a huge unexploited resource. • Mesopelagics have multiple usage areas such as fish feed and direct human consumption. • Knowledge generation of the mesopelagic zone is vital before a fishery can occur. [ABSTRACT FROM AUTHOR]

Calanus finmarchicus, like many other copepods in the family Calanidae, can enter into a facultative diapause during the last juvenile phase (fifth copepodid, C5) to enable survival during unfavorable periods. Diapause is essential to the persistence of Calanus populations and profoundly impacts energy flow within oceanic ecosystems, yet regulation of diapause is not understood in these animals. Transcriptional profiling has begun to provide insight into metabolic changes occurring as C. finmarchicus prepares for and enters into diapause or skips diapause to prepare for the terminal molt. In particular, components of the glycolysis, pentose phosphate and lipid synthesis pathways are upregulated early in the C5 stage when lipid stores are low. Currently, our ability to identify metabolic patterns is limited by the incomplete functional annotation of the C. finmarchicus transcriptome. Such limitations are widespread among studies of non-model organisms and addressing them should be a priority for future research. In addition, integrating the results across multiple emerging complementary transcriptomic studies will provide a more complete picture of copepod physiology than isolated studies. Ultimately, identifying molecular markers of copepod physiology could enable robust identification of animals preparing to enter into diapause and ultimately lead to a greatly improved understanding of diapause regulation. [ABSTRACT FROM AUTHOR]

Climate change is affecting numerous species worldwide, including dominant and important copepods of the genus Calanus . Despite the growing body of studies that examine effects of climate change stressors on Calanus species, comparative intraspecific studies are lacking. Importantly, acclimatization and genetic adaptation can modify the stress response, thus leading to a differential response of separated populations to the same stressor. The molecular and physiological responses of a C. finmarchicus population from the mid-Atlantic, with an in situ temperature of 8.5 °C, were investigated under experimental thermal conditions of 0 °C, 5 °C, 10 °C, 15 °C, and 20 °C for durations of 3 h and 6 days. This experimental set-up mirrored previously published experiments conducted on C. finmarchicus at the northern limit of its distribution allowing a comparison between two populations. The greatest physiological response, assessed as fecal pellet production, was seen after 3 h exposure at 10 °C and 15 °C, and after 6 days exposure at 5 °C, 10 °C and 15 °C. Molecular response was assessed by the change in expression of 5 selected genes: hsp70 _ 2 , dnaja1 , nap 1l1 , rps11 , and gdh . Only two out of the five genes ( gdh and nap1l1 ) showed significant up-regulation with increased temperature and duration of exposure. These findings differ from the results obtained in the northern population where all 5 genes were differently expressed. Overall, the results suggest population-specific response to temperature in C. finmarchicus , however determining the source of such variation (genetic adaptation or acclimatization) requires more detailed studies. [ABSTRACT FROM AUTHOR]

Several abstracts related to lipid technology are presented including Canola engineered with microalgal polyketide synthase-like system, wax ester rich oil from Calanus finmarchicus, and the level of omega-3 index in people with schizophrenia and depression.

Diapause plays a key role in the life cycle of high latitude zooplankton. During diapause, animals avoid starving in winter by living in deep waters where metabolism is lower and met by lipid reserves. Global warming is therefore expected to shorten the maximum potential diapause duration by increasing metabolic rates and by reducing body size and lipid reserves. This will alter the phenology of zooplankton, impact higher trophic levels and disrupt biological carbon pumps. Here, we project the impacts of climate change on the key North Atlantic copepod Calanus finmarchicus under IPCC RCP 8.5. Potential diapause duration is modelled in relation to body size and overwintering temperature. The projections show pronounced geographic variations. Potential diapause duration reduces by more than 30% in the Western Atlantic, whereas in the key overwintering centre of the Norwegian Sea it changes only marginally. Surface temperature rises, which reduce body size and lipid reserves, will have a similar impact to deep-water changes on diapause in many regions. Because deep-water warming lags that at the surface, animals in the Labrador Sea could offset warming impacts by diapausing in deeper waters. However, the ability to control diapause depth may be limited. [ABSTRACT FROM AUTHOR]

Marine oils are rich in polyunsaturated fatty acids (PUFAs), including docosahexaenoic and eicosapentaenoic acid. These PUFAs are associated with health benefits and additional sustainable sources of marine oils are desirable. One of the source organisms is Calanus finmarchicus , a copepod endemic to the North Atlantic. PUFAs in the lipid fraction of this organism are largely in the form of wax esters. To assess the safety of these wax esters as a source of PUFAs, a randomized, double-blinded, placebo-controlled clinical trial was conducted whereby 64 subjects consumed 2 g Calanus oil in capsule form daily for a period of one year. A group of 53 subjects consumed placebo capsules. At baseline, 6-, and 12-months, series of evaluations were conducted, including: vital signs, clinical chemistry and hematological evaluations, and adverse event reporting. Food intake and physical exercise were controlled by means of a questionnaire. There were no effects on Calanus oil treatment on any of the safety parameters measured. A slight increase in the incidence of eczema was reported in the Calanus oil group, but the response was minor in nature, not statistically significant after controlling for multiple comparisons, and could not be attributed to treatment. [ABSTRACT FROM AUTHOR]

Zooplankton abundance and community structures were studied in three west Spitsbergen fjords at the beginning of the warm phase, which seem to have entered in 2006. Sampling was conducted in summer 2007 at stations distributed along transects in Hornsund, Isfjorden and Kongsfjorden. Variations in zooplankton standing stocks and community structures (assessing taxonomic diversity and zoogeographical affiliations) were analysed in relation to the environmental variables using multivariate techniques. The hydrographic conditions in Hornsund were influenced by the cold Arctic Water, whereas those in Isfjorden and especially in Kongsfjorden were, to a greater extent, under the influence of the warm Atlantic Water. High abundances of both meroplankton and holoplankton organisms were observed in Kongsfjorden, with high contributions of boreal and ubiquitous species ( Calanus finmarchicus and Oithona similis, respectively). In Hornsund at the same time, the zooplankton consisted mainly of boreo-Arctic and Arctic species, the abundances of which were comparable along the West Spitsbergen Shelf. Our results indicate that the difference in hydrography had measurable effects on the zooplankton community in the study area. Furthermore, by comparing regions of contrasting oceanographic conditions, we present evidence as to how the zooplankton structure will change in the Arctic ecosystems if the warming trends continue to operate with the same dynamics. The advection of Atlantic waters to the Arctic seas may lead to changes in zooplankton structure, with increased abundance and contributions of boreal and small ubiquitous species. The 'warmer Arctic fjords' scenarios may also induce more rapid development of both holoplankton and meroplankton populations and, consequently, modify the trophic interactions in plankton communities. [ABSTRACT FROM AUTHOR]

Arctic species of Calanus are critical to energy transfer between higher and lower trophic levels and their relative abundance, and lipid content is influenced by the alternation of cold and warm years. All three species of Calanus were collected during different periods in Kongsfjorden (Svalbard, 79°N) and adjacent shelf during the abnormally warm year of 2006. Lipid composition and fatty acid structure of individual lipid classes were examined in relation with population structure. Wax esters dominated the neutral lipid fraction. Phosphatidylcholine (PC) dominated the structural lipids followed by phosphatidylethanolamine (PE). PC/PE ratios of 3-6 suggested an increase in PC proportions compared to earlier studies. Depending on the time scale, fatty acids of wax esters illustrated either trophic differences between fjord and offshore conditions for C. hyperboreus and C. finmarchicus or trophic differences related to seasonality for C. glacialis. Similarly, seasonality and trophic conditions controlled the changes in fatty acids of triglycerides, but de novo synthesis of long-chain monoenes suggested energy optimization to cope with immediate metabolic needs. Polar lipids fatty acid composition was species specific and on the long-term (comparison with data from the past decade) composition appears related to changes in trophic environment. Fatty acid composition of PC and PE indicated relative dominance of 20:5n-3 in PC and 22:6n-3 in PE for all three species. The combination of PE and PC acyl chain and phospholipid head group restructuring indicates an inter-annual variability and suggests that membrane lipids are the most likely candidate to evaluate adaptive changes in Arctic copepods to hydrothermal regime. [ABSTRACT FROM AUTHOR]

Near-bottom zooplankton communities have rarely been studied despite numerous reports of high zooplankton concentrations, probably due to methodological constraints. In Kongsfjorden, Svalbard, the near-bottom layer was studied for the first time by combining daytime deployments of a remotely operated vehicle (ROV), the optical zooplankton sensor moored on-sight key species investigation (MOKI), and Tucker trawl sampling. ROV data from the fjord entrance and the inner fjord showed high near-bottom abundances of euphausiids with a mean concentration of 17.3 ± 3.5 n × 100 m. With the MOKI system, we observed varying numbers of euphausiids, amphipods, chaetognaths, and copepods on the seafloor at six stations. Light-induced zooplankton swarms reached densities in the order of 90,000 (euphausiids), 120,000 (amphipods), and 470,000 ind m (chaetognaths), whereas older copepodids of Calanus hyperboreus and C. glacialis did not respond to light. They were abundant at the seafloor and 5 and 15 m above and showed maximum abundance of 65,000 ind m. Tucker trawl data provided an overview of the seasonal vertical distribution of euphausiids. The most abundant species Thysanoessa inermis reached near-bottom concentrations of 270 ind m. Regional distribution was neither related to depth nor to location in the fjord. The taxa observed were all part of the pelagic community. Our observations suggest the presence of near-bottom macrozooplankton also in other regions and challenge the current view of bentho-pelagic coupling. Neglecting this community may cause severe underestimates of the stock of pelagic zooplankton, especially predatory species, which link secondary production with higher trophic levels. [ABSTRACT FROM AUTHOR]

We conducted a year-round mesozooplankton study in the Arctic Kongsfjord from August 1998 until July 1999 to investigate seasonal abundance and vertical as well as stage distributions of the prevalent taxa. It is the first investigation in Kongsfjord that covers the Arctic winter season and provides reasonable estimates also of small-sized copepod species. Abundant smaller copepods comprised Oithona similis, Pseudocalanus minutus, Microcalanus spp., Triconia borealis and Acartia longiremis. Among the larger copepods, Calanus finmarchicus, C. glacialis, C. hyperboreus and Metridia longa dominated. The thecosome pteropod Limacina helicina was also an important component. Abundance maxima occurred in November (988,669 ind. m) with one to two orders of magnitude higher numbers as compared to all other months (39,832-200,067 ind. m). The summers of 1998 and 1999 were characterized by intrusions of Atlantic water, but the community was not entirely dominated by advected boreal species. During winter, the majority of the mesozooplankton occurred below 100 m. Advection is the most likely reason for the accumulation of zooplankton at depth in winter, but local production may also contribute to high overwintering numbers. Much lower abundances of most species in spring suggest high winter mortality and emphasize the importance of sufficient reproductive success during the previous summer to ensure enough winter survivors as seed stock for the coming reproductive season. This study was conducted prior to the recent warming trend in the Arctic. Therefore, it provides valuable baseline data and allows comparing present and future states of the zooplankton community in Kongsfjord. [ABSTRACT FROM AUTHOR]

Calanus finmarchicus , an abundant calanoid copepod in the North Atlantic Ocean, is both a major grazer on phytoplankton and an important forage species for invertebrate and vertebrate predators. One component of the life history of C. finmarchicus is the overwintering dormancy of sub-adults, a feature key for the annual recruitment of this species in early spring. While little is known about the control of dormancy in C. finmarchicus , one hypothesis is that it is an insect-like diapause, where the endocrine system is a key regulator. One group of hormones implicated in the control of insect diapause is the insulin-like peptides (ILPs). Here, C. finmarchicus transcriptomic data were used to predict ILP signaling pathway proteins. Four ILP precursors were identified, each possessing a distinct A- and B-chain peptide; these peptides are predicted to form bioactive heterodimers via inter-chain disulfide bridging. Two ILP receptors, which likely represent splice variants of a common gene, were identified. Three insulin-degrading enzymes were also discovered, as were proteins encoding the transcription factor FOXO, a downstream target of ILP that has been implicated in the regulation of insect diapause, and insulin receptor substrate, a protein putatively linking the ILP receptor and FOXO. RNA-Seq data suggest that some C. finmarchicus insulin pathway transcripts are differentially expressed across development. As in insects, the ILP signaling system may be involved in controlling C. finmarchicus ’ organism-environment interactions ( e.g. , regulation of seasonal sub-adult diapause), a hypothesis that can now be investigated using these data. [ABSTRACT FROM AUTHOR]

Deep convection in the Labrador and Irminger Seas inflates the cold and low-saline subpolar gyre, which is a rich nutrient and zooplankton source for the surrounding warmer waters of subtropical origin. The zooplankton abundances on the south Iceland shelf show characteristic sub-decadal variability, which closely reflect the oceanic abundances of the ecologically most important zooplankton species – Calanus finmarchicus. Much higher abundances of this species are observed during years when the winter mixed layer depths (MLD) in the Labrador-Irminger Sea, and over the Reykjanes Ridge are deep. Furthermore, a tight relationship is identified between on-shelf zooplankton abundances and lateral shifts of the biologically productive subarctic front southwest of Iceland. Thus, we suggest that northeastward expansion of the subpolar gyre results in biologically productive periods in the waters southwest of Iceland – both oceanic and on the shelf. In addition to local atmospheric forcing, we find that the MLD and frontal position are also impacted by remote heat losses and convection in the Labrador Sea, through northward advection of unstable mode waters. The sub-decadal oceanic and on-shelf biological production peaks are possibly predictable by half a year (local winter convection to subsequent summer production), and the advective time-lag from the Labrador Sea might induce an even longer predictability horizon (up to 1.5 years). [ABSTRACT FROM AUTHOR]

How the distribution of Calanus finmarchicus and its potential northward expansion will be affected by climate changes depends on the mechanisms and processes constraining their reproduction, recruitment and survival. Here we present measurements of C. finmarchicus egg production rates during the spring bloom in 2008, 2010 and 2011 in Disko Bay, West Greenland and validate four independently derived metabolic models to predict egg production rates. The spring bloom in 2008 was short and intense and supported lower cumulated specific egg production of C. finmarchicus than the longer blooms with lower peak biomass in 2011 and 2012. The models predicted different timing of initialization and development of egg production rates based on phytoplankton biomass and temperature and model performance varied from 'poor' to 'very good'. Phytoplankton biomass controlled the changes in egg production rates during the spring in Disko Bay, while the low temperature in the Bay explained why the egg production rate here is much lower than at more southerly localities despite high food concentrations. This study suggests that an increase in magnitude of the Arctic phytoplankton spring bloom will not result in increased copepod egg production, whereas a longer bloom and increasing temperatures will. [ABSTRACT FROM AUTHOR]

Calanus finmarchicus is an important zooplankton species in the Norwegian Sea, as a dominant food organism for pelagic fish larvae, and a potentially large source of marine lipids and proteins. Its position in the marine food web also makes it an important model species in assessing the risk posed by oil spills in the Norwegian and Arctic Seas. In this study, an Eulerian population model for C.finmarchicus , coupled to the physical and ecological model SINMOD, is presented. The model includes the full life cycle of C. finmarchicus with a representation of all developmental stages. The model has been validated against field measurements made in different areas of the Norwegian Sea in 1997 and 1998. The model displays geographical and temporal distributions of development stages that is in line with observed patterns. When comparing time series for selected regions, we see a high degree of variability both in the field samples and model output. On average, the model deviations are near half of the summed variability of the field data and model estimates. The model has applications within assessment of ecological production, and the potential for harvesting in the Norwegian and Arctic Seas, but in combination with other models, also for the assessment of ecological effects of oil spills and other types of pollution. [ABSTRACT FROM AUTHOR]

In the current study differences were evaluated between a complex 3D multistage population model (SINMOD) and a simpler consumer-resource population model for estimating the effects of crude oil on the marine copepod Calanus finmarchicus . The SINTEF OSCAR model was used to simulate hypothetical oil spills in the Lofoten area in 1995, 1997, and 2001. Both population models simulated a negligible effect of crude oil on the Calanus’ biomass when assuming low species sensitivity. The simple model estimated a larger effect on the biomass (up to a 100% decline) compared to the complex model (maximum decline of 60–80%) at high species sensitivity to crude oil. These differences may be related to the inclusion of copepod advection in the complex model. Our study showed that if little data is available to parameterize a model, or if computational resources are scarce, the simple model could be used for risk screening. Nevertheless, the possibility of including a dilution factor for time-varying biomass should be examined to improve the estimations of the simple model. The complex model should be used for a more in depth risk analysis, as it includes physical processes such as the drift of organisms and differentiation between developmental stages. [ABSTRACT FROM AUTHOR]

The effect of the dinoflagellate, Alexandrium fundyense, on relative expression of glutathione S-transferase (GST) transcripts was examined in the copepod Calanus finmarchicus. Adult females were fed for 5-days on one of three experimental diets: control (100% Rhodomonas spp.), low dose of A. fundyense (25% by volume, 75% Rhodomonas spp.), and high dose (100% A. fundyense). Relative expression of three GST genes was measured using RT-qPCR on days 0.5, 1, 2 and 5 in two independent experiments. Differential regulation was found for the Delta and the Sigma GSTs between 0.5 to 2 days, but not on day 5 in both experiments. The third GST, a microsomal, was not differentially expressed in either treatment or day. RT-qPCR results from the two experiments were similar, even though experimental females were collected from the Gulf of Maine on different dates and their reproductive output differed. In the second experiment, expression of 39 GSTs was determined on days 2 and 5 using RNA-Seq. Global gene expression analyses agreed with the RT-qPCR results. Furthermore, the RNA-Seq measurements indicated that only four GSTs were differentially expressed under the experimental conditions, and the response was small in amplitude. In summary, the A. fundyense diet led to a rapid and transient response in C. finmarchicus in three cytosolic GSTs, while a fourth GST (Omega I) was significantly up-regulated on day 5. Although there was some regulation of GSTs in response the toxic dinoflagellate, the tolerance to A. fundyense by C. finmarchicus is not dependent on the long-term up-regulation of specific GSTs. [ABSTRACT FROM AUTHOR]

Marine populations are controlled by a series of drivers, pertaining to both the physical environment and the biological environment (trophic predator-prey interactions). There is heated debate over drivers, especially when trying to understand the causes of major ecosystem events termed regime shifts. In this work, we have researched and developed a novel methodology based on Genetic Programming (GP) for distinguishing which drivers can influence species abundance. This methodology benefits of having no a priori assumptions either on the ecological parameters used or on the underlying mathematical relationships among them. We have validated this methodology applying it to the North Sea pelagic ecosystem. We use the target species Calanus finmarchicus, a key copepod in temperate and subarctic ecosystems, along with 86 biological, hydrographical and climatic time series, ranging from local water nutrients and fish predation, to large scale climate pressure patterns. The chosen study area is the central North Sea, from 1972 to 2011, during which period there was an ecological regime shift. The GP based analysis identified 3 likely drivers of C. finmarchicus abundance, which highlights the importance of considering both physical and trophic drivers: temperature, North Sea circulation (net flow into the North Atlantic), and predation (herring). No large scale climate patterns were selected, suggesting that when there is availability of both data types, local drivers are more important. The results produced by the GP based procedure are consistent with the literature published to date, and validate the use of GP for interpreting species dynamics. We propose that this methodology holds promises for the highly non-linear field of ecology. [ABSTRACT FROM AUTHOR]

In the Barents Sea region new petroleum fields are discovered yearly and extraction of petroleum products is expected to increase in the upcoming years. Despite enhanced technology and stricter governmental legislation, establishment of the petroleum industry in the Barents Sea may potentially introduce a new source of contamination to the area, as some discharges of produced water will be allowed. Whether the presence of produced water poses a risk to the Arctic marine life remains to be investigated. The aim of this study was to examine effects of exposure to several compounds found in produced water—a mixture of selected organic compounds (APW), radium-226 (226Ra), barium (Ba), and a scale inhibitor—on the copepod speciesCalanus finmarchicus. Experiments were performed using exposure concentrations at realistic levels based on those detected in the vicinity of known discharge points. The influence of lethal and sublethal effects on early life stages was determined and significantly lower survival in the APW exposure groups was found. In the Ba treatment the life stage development did not proceed to the same advanced stages as observed in the control (filtered sea water). The scale inhibitor and226Ra treatments showed no significant difference from control. In addition, adult females were exposed to APW,226Ra, and a mixture of the two. Both individual-level effects (egg production and feeding) and molecular-level effects (gene expression) were assessed. On the individual level endpoints, only treatments including APW produced an effect compared to control. However, on the molecular level the possibility that also226Ra induced toxicologically relevant effects cannot be ruled out. [ABSTRACT FROM PUBLISHER]

In this investigation, acute toxicity data were used from two previously reported studies where cold-water copepods were exposed to mechanically dispersed (MD) and chemically (CD) dispersed oil. In one of these studies, concentration-dependent mortality was observed, whereas no apparent relationship between exposure concentration and mortality was found in the other. The only marked difference between the studies is that copepods in the first experiment displayed a lower lipid sac volume (on average) than in the second one. In this study additional biometric data on lipid content were utilized and observed effects and toxicokinetics modeling applied in order to investigate whether differences in sensitivity between copepod cohorts might be explained by differences in lipid content. Results suggest that although a considerable lipid sac might retard toxicokinetics, the observed differences in lipid volume are not sufficient to explain differences in toxicity. Further, there are no apparent indications that acute toxic stress leads to lipid depletion, or that acute increased mortality rate selectively affects lipid-poor individuals. It is conceivable that other potential explanations exist, but the causal relationship between lipid content and increased mortality frequency remains elusive. [ABSTRACT FROM PUBLISHER]

Mackerel ( Scomber scombrus) is one of the ecologically and economically most important fish species in the Atlantic. Its recruitment has, for unknown reasons, been exceptional from 1998 to 2012. The majority (75%) of the survivors in the first winter were found north of an oceanographic division at approximately 52°N, despite the fact that mackerel spawns over a wide range of latitudes. Multivariate time series modelling of survivor abundance in the north revealed a significant correlation with the abundance of copepodites (stage I-IV) of Calanus sp. in the spawning season (April to June). The copepodites were a mix of C. helgolandicus (dominating) and C. finmarchicus. The growth of mackerel larvae is known to be positively related to the availability of nauplii and copopodites of preferred prey species, namely, large calanoid copepod species such as Calanus. The statistical relationship between mackerel survivors and abundance of Calanus, therefore, most likely, reflected a causal relationship: high availability of Calanus probably reduced starvation, stage-specific predation and cannibalism (owing to prey switching). The effects of other abundant, but less preferred zooplankton taxa, ( Acartia sp., Branchiopoda spp. and Echinodermata spp. larvae), as well as stock size, temperature and wind-induced turbulence were not found to be significant. However, stock size was retained in the final model because of a significant interaction with Calanus in oceanic areas west of the North European continental shelf. This was suggested to be a consequence of a density driven expansion of the spawning area that increased the overlap between early life stages of mackerel and food ( Calanus) in new areas. [ABSTRACT FROM AUTHOR]

The southwestern Norwegian Sea is characterized by an inflow of warm and saline Atlantic water from the southwest and cold and less saline East IcelandicWater (EIW), of Arctic origin, from the northwest. These two water masses meet and form the Iceland-Faroe Front (IFF). In this region, the copepod Calanus finmarchicus plays a key role in the pelagic ecosystem. Time-series of C. finmarchicus and Calanus hyperboreus in May and September, extending back to the early 1990s,were studied in relation to phytoplankton bloom dynamics and hydrography. The main reproductive period of C. finmarchicus started consistently earlier south of the IFF, resulting in different life cycles and stage compositions in the twowater masses. In 2003, a sudden shift occurred north of the IFF, resulting in a similar phenology pattern to south of the IFF. Before this, only one generation of C. finmarchicus was produced in the Arctic water, but the earlier reproduction enabled the species to produce two generations after 2003. Simultaneously, C. hyperboreus, an expatriate in the EIW, largely disappeared. Food availability is unlikely the reason for the phenological differences observed across the front, as the typical pattern of the phytoplankton spring bloom showed an earlier onset north of the IFF. Temperature and salinity peaked at record high values in 2003 and 2004, and therefore possible links to oceanography are discussed. The dominant role of Calanus spp. and the potential linkages to water mass exchanges may herald strong effects on the ecosystem and pelagic fish in this subpolar Atlantic region under expected climate change. [ABSTRACT FROM AUTHOR]

This study investigates whether feeding on the domoic acid (DA)-producing diatom Pseudo-nitzschia seriata affects the faecal pellet (FP) production (proxy for grazing) and fecundity of Calanus finmarchicus and Calanus glacialis. Female copepods were fed a saturating concentration of food (400 μg C L-1) in two combinations (i) natural phytoplankton spiked with 50% P. seriata and (ii) only the non-toxic phytoplankton community. The copepods ingested DA, as illustrated by DA accumulation in their FPs, and transferred a share of DA to their eggs and body tissue. DA was mainly excreted through FPs in both species. Compared with C. finmarchicus, C. glacialis accumulated substantially more DA in its body tissue. For both species, egg production and hatching success were unaffected by grazing on the toxic diatom. This suggests that feeding on DA-containing P. seriata does not affect copepod fecundity, despite increasing DA concentrations of P. seriata during the experiment. [ABSTRACT FROM AUTHOR]

Mortality is notoriously difficult to estimate for zooplankton populations in the open ocean due to the confounding effect of advection. The vertical life table (VLT) approach is commonly used, but has been shown to be sensitive to both spatial and temporal trends in recruitment. Here, we estimate mortality rates of Calanus finmarchicus copepodites from spatiotemporally resolved data from the highly advective Norwegian Sea-Barents Sea in spring and summer. We apply both the VLT and a statistical regression approach (SRA), specifically taking into account the effects of advection and spatiotemporal trends in recruitment. Testing the two methods on a simulated dataset shows that the SRA performs better than the most commonly used version of the VLT when trends in recruitment are present. Overall, the SRA appears to be a robust method for spatiotemporally resolved survey data influenced by advection and spatiotemporal trends in recruitment. The estimated mortality rates are relatively low (0.03-0.07 d-1) and indicate increased mortality for the oldest stage pair (copepodites CIV-CV) compared to earlier stages. The results are discussed in the light of previous mortality rate estimates for C. finmarchicus. [ABSTRACT FROM AUTHOR]

High Calanus finmarchicus abundances were recorded in wintertime in Vestfjorden, close to the main cod breeding grounds off Lofoten and Vesterålen, northern Norway. The mean abundance for locations with water depth >500 m was ~37000 ind. m-2 (range: 26700-49000 ind. m-2). To our knowledge, this is the first report of massive overwintering of C. finmarchicus on the Norwegian shelf. Because of the observed size and location of this population, we argue that local overwintering on the northern Norwegian shelf can contribute significantly to sustain a C. finmarchicus population on the shelf during the period of first feeding for cod larvae. This is supported by a particle tracking model. [ABSTRACT FROM AUTHOR]

Acute toxicity differs between species but also varies within a species. Important intraspecific factors are the exposure duration and properties of the animal such as life stage, sex, and physiological status. In the present study, the acute toxicity of water-soluble fractions (WSFs) from fresh and artificially weathered oil was followed over time in different life stages of the calanoid copepod Calanus finmarchicus, including adult males and females. The life stages differ in size but also in lipid content and physiology. To meaningfully compare the sensitivity of the different stages, the authors fitted a toxicokinetic-toxicodynamic (TKTD) model from the framework of the General Unified Threshold Model of Survival (GUTS) to the mortality patterns over time. The oil WSFs could not be treated as single compounds: the rapid effect at high doses could not be reconciled with the slow effect at low doses. Treating the oil as a mixture of 2 component blocks could, however, capture these patterns satisfactorily. Even though the early life stages of animals are generally considered to be the most vulnerable, the adult males of C. finmarchicus turned out to be most sensitive, followed by the early copepodites. Naupliar larvae were equally susceptible to oil toxicity as late copepodites and adult females. The relationship between the GUTS model parameters and the physiological traits for the different life stages remains, however, unclear. Environ Toxicol Chem 2016;35:728-735. © 2015 SETAC [ABSTRACT FROM AUTHOR]

Understanding how climate influences ecosystems is complicated by the many correlated and interrelated impacting factors. Here we quantify climate effects on Calanus finmarchicus in the northeastern Norwegian Sea and southwestern Barents Sea. By combining oceanographic drift models and statistical analyses of field data from 1959 to 1993 and investigating effects across trophic levels, we are able to elucidate pathways by which climate influences zooplankton. The results show that both chlorophyll biomass in spring and C. finmarchicus biomass in summer relate positively to a combination of shallow mixed layer depth and increased wind in spring, suggesting that C. finmarchicus biomass in summer is influenced by bottom-up effects of food availability. Furthermore, spatially resolved C. finmarchicus biomass in summer is linked to favorable transport fromwarmer, core areas to the south. However, increased mean temperature in spring does not lead to increased C. finmarchicus biomass in summer. Rather, spring biomass is generally higher, but population growth from spring to summer is lower, after a warm compared with a cold spring. Our study illustrates how improved understanding of climate effects can be obtained when different datasets and different methods are combined in a unified approach. [ABSTRACT FROM AUTHOR]

The Subarctic copepod, Calanus finmarchicus, is an ecologically critical foundation species throughout the North Atlantic Ocean. Any change in the abundance and distribution of C. finmarchicus would have profound effects on North Atlantic pelagic ecosystems and the services that they support, particularly on the coastal shelves located at the southern margins of the species' range. We tested the hypothesis that the physiological rates and processes of C. finmarchicus, determining its vital rates, are unaffected by increases in CO2 concentration predicted to occur in the surface waters of the ocean during the next 100 years. We reared C. finmarchicus from eggs to adults at a control (580 µatm, the ambient concentration at the laboratory's seawater intake) and at predicted mid-range (1200 µatm) and high (1900 µatm) pCO2. There was no significant effect of pCO2 on development times, lipid accumulation, feeding rate, or metabolic rate. Small but significant treatment effects were found in body length and mass (in terms of dry, carbon and nitrogen mass), notably a somewhat larger body size at the mid-pCO2 treatment; that is, a putatively beneficial effect. Based on these results, and a review of other studies of Calanus, we conclude that the present parameterizations of vital rates in models of C. finmarchicus population dynamics, used to generate scenarios of abundance and distribution of this species under future conditions, do not require an "ocean acidification effect" adjustment. A review of research on planktonic copepods indicates that, with only a few exceptions, impacts of increased CO2 are small at the levels predicted to occur during the next century. [ABSTRACT FROM AUTHOR]

It is currently under debate whether organisms that regulate their acid-base status under environmental hypercapnia demand additional energy. This could impair animal fitness, but might be compensated for via increased ingestion rates when food is available. No data are yet available for dominant Calanus spp. from boreal and Arctic waters. To fill this gap, we incubated Calanus glacialis at 390, 1120, and 3000 µatm for 16 d with Thalassiosira weissflogii (diatom) as food source on-board RV Polarstern in Fram Strait in 2012. Every 4 d copepods were subsampled from all CO2 treatments and clearance and ingestion rates were determined. During the SOPRAN mesocosm experiment in Bergen, Norway, 2011, we weekly collected Calanus finmarchicus from mesocosms initially adjusted to 390 and 3000 µatm CO2 and measured grazing at low and high pCO2. In addition, copepods were deep frozen for body mass analyses. Elevated pCO2 did not directly affect grazing activities and body mass, suggesting that the copepods did not have additional energy demands for coping with acidification, neither during long-term exposure nor after immediate changes in pCO2. Shifts in seawater pH thus do not seem to challenge these copepod species. [ABSTRACT FROM AUTHOR]

Early life stages of fish spawned on the Norwegian continental shelf have long been suggested to depend on eggs and nauplii from the crustacean zooplankton Calanus finmarchicus for survival. Calanus finmarchicus overwinters in the deep basins of the Norwegian Sea, and gravid females must be advected onto the shelf prior to spawning if eggs and nauplii larvae are to serve as food for fish larvae. In this study, cross-shelf advection of C. finmarchicus is simulated over 52 years (1960-2011) using a numerical ocean model coupled with an individual-based model. The results suggest that cross-shelf transport of C. finmarchicus is surprisingly stable across years and that transport is particularly concentrated immediately upstream of the two major spawning areas for the Northeast Arctic cod (Gadus morhua) and the Norwegian spring-spawning herring (Clupea harengus), namely Lofoten and Møre, respectively. Two large topographical features, the Træna Trough and the Norwegian Trench, appear to be funnelling C. finmarchicus onto the shelf in these two areas. This could suggest that the fish spawning grounds outside Møre and Lofoten are, in part, maintained owing to stable interannual food supply in spring. [ABSTRACT FROM AUTHOR]

Inshore and offshore waters of the Gulf of Maine (USA) have spring/summer harmful algal blooms (HABs) of the toxic dinoflagellate Alexandrium fundyense , which is responsible for paralytic shellfish poisoning (PSP) in humans. The calanoid copepod Calanus finmarchicus co-occurs with A. fundyense during the seasonal blooms. At that time, C. finmarchicus population abundances are high, dominated by immature copepods preparing for diapause, and by actively-reproducing adults. High survival has been reported for copepods exposed to toxic A. fundyense , but little is known about possible sublethal effects. In this study, C. finmarchicus adult females were fed either a control diet of non-toxic Rhodomonas spp. or one of two diets containing either low dose (LD) or high dose (HD) levels (50 and 200 cells mL −1 , respectively) of toxic A. fundyense for a total of 7 days in two independent experiments. As expected, ingestion of the dinoflagellate had no effect on copepod survival and grazing activity. However, significant reductions of egg production and egg viability were observed in C. finmarchicus females fed on either experimental diet . After the 7-day experiment, total nauplius production by females on the LD and HD diets was reduced by 35% to 75% compared to the control females. These results suggest that blooms of A. fundyense in the Gulf of Maine may be an environmental challenge for C. finmarchicus populations, with a potential negative effect on copepod recruitment. [ABSTRACT FROM AUTHOR]

The Continuous Plankton Recorder (CPR) survey has collected plankton since 1958 in the North Atlantic Ocean and its adjacent seas. Among all species recorded by the CPR, Calanus finmarchicus has probably been the most investigated species because of its ecological importance for the temperate and subpolar regions of the North Atlantic Ocean. However, abundances of C. finmarchicus assessed from the CPR survey have been rarely compared to more traditional sampling methodologies. In this study, we examine and compare spatial (surface and vertical) and temporal (diel and seasonal) patterns in the abundance of C. finmarchicus with another sampling technique in the gulf of Maine. Our results provide evidence that the CPR survey not only gives internally consistent time series of C. finmarchicus , but also an accurate representation of both spatial (surface and vertical) and temporal (diel and seasonal) patterns. [ABSTRACT FROM AUTHOR]