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Investigations of genetic stock structure sometimes reveal a mismatch between management units and biological units. In Scandinavian waters, Norway lobster (Nephrops norvegicus) is divided into two management units (the Skagerrak–Kattegat and the Norwegian Deep). We have tested the population genetic structure of Nephrops within this region using microsatellite DNA markers, and compared the structure with the present management units. Our study suggests no population genetic structure of Nephrops within the Skagerrak, Kattegat, and Norwegian Deep region, whereas a shallow genetic structure was detected on a larger geographical scale when comparing outgroup samples from Scotland and Iceland. We found indications of sex-biased dispersal as the overall genetic differences were larger for females. Ocean current patterns suggest that Nephrops stocks in the region may be connected by larval drift. The two areas differ in fishing pressure, monitoring, assessment, and regulations, which is an argument for maintaining the present two-areas management regime despite the evidence for one biological population.

Environmental DNA (eDNA) has gained popularity as a tool for ecosystem biomonitoring and biodiversity assessment. Although much progress has been made regarding laboratory and fieldwork protocols, the issue of sampling efficiency requires further investigation, particularly in three-dimensional marine systems. This study focuses on fish community composition in marine ecosystems and aims to analyze the efficiency of sampling design given the sampling effort for distinguishing between different communities. We sampled three fjords in Northern Norway, taking samples along fjord transects and at three different depths, and amplified a fragment of the mitochondrial 12S rRNA gene of bony fishes using the MiFish primers. We evaluated the effect of (i) the number of sampling stations, (ii) samples' spatial distribution, and (iii) the data treatment approach (presence/absence versus semiquantitative) for maximizing the efficiency of eDNA metabarcoding sampling when inferring differences of fish community compositions between fjords. We found that the manner of data treatment strongly affected the minimum number of sampling stations required to detect differences among communities; because the semiquantitative approach retained some information about abundance of the underlying reads, it was the most efficient. Furthermore, we found little-to-no difference of fish communities in samples from intermediate depths when comparing vertical fish communities. Lastly, we found that the differences between fish communities at the surface were the highest across the horizontal distance and overall, samples ~30 km apart showed the highest variation in the horizontal distribution. Boosting sampling efficiency (reducing sampling effort without compromising ecological inferences) can significantly contribute to enhanced biodiversity management and efficient biomonitoring plans.

Identifying how physical and biotic factors shape genetic connectivity among populations in time and space is essential to our understanding of the evolutionary trajectory as well as the management of marine species. Atlantic cod is a widespread and commercially important marine species displaying several ecotypes with different life history strategies. Using three sets of SNPs: neutral, informative, and genome-inversion linked, we studied population genetic structure of ~2500 coastal Atlantic cod (CC) from 40 locations along Norway's 2500 km coastline, including nine fjords. We observed: (1) a genetic cline, suggesting a mechanism of isolation by distance, characterized by a declining FST between CC and North East Arctic Cod (NEAC—genetically distinct migratory ecotype) with increasing latitude, (2) that in the north, samples of CC from outer-fjord areas were genetically more similar to NEAC than were samples of CC from their corresponding inner-fjord areas, (3) greater population genetic differentiation among CC sampled from outer-fjord areas along the coast, than among CC sampled from their corresponding inner-fjord areas, (4) genetic differentiation among samples of CC from both within and among fjords. Collectively, these results permit us to draw two main conclusions. First, that differences in the relative presence of the genetically highly distinct, migratory ecotype NEAC, declining from north to south and from outer to inner fjord, plays the major role in driving population genetic structure of the Norwegian CC. Second, that there is limited connectivity between CC from different fjords. These results suggest that the current management units implemented for this species in Norway should be divided into smaller entities. Furthermore, the situation where introgression from one ecotype drives population genetic structure of another, as is the case here, may exist in other species and geographical regions, thus creating additional challenges for sustainable fisheries management. publishedVersion

Marine fisheries are often allocated to stocks that reflect pragmatic considerations and may not represent the species’ spatial population structure, increasing the risk of mismanagement and unsustainable harvesting. Here we compile mark–recapture data collected across the North Atlantic to gain insight into the spatial population structure of Greenland halibut (Reinhardtius hippoglossoides), an issue that has been unresolved for decades. The dataset contains 168130 fish tagged from 1952 to 2021, with 5466 (3.3%) recaptured individuals. Our results indicate that fish tagged at

Blue ling is a deep-water species that has been severely fished upon into coastal and offshore fisheries since the early 1970s, thus causing the collapse of the populations in the last two decades. Genetic information is scarce in this species, and molecular markers are therefore needed to provide advice both for management and for rebuilding the stocks. A suite of 103 SNP markers was identified from ddRAD sequencing data. From those, 81 were organized in three multiplex reactions, and tested on 150 individuals from three different sampling locations. Good-quality amplification products were successfully obtained from 70 of the markers. All SNP loci were biallelic, with averaged He per locus ranging between 0.101 and 0.500.

Coastal Atlantic cod (Gadus morhua) in the Northeast Atlantic has seen a continuous decline since the industrialization of the coastal fishery, and management needs to address the spatial and temporal complexities of coexisting cod stocks. Toward that end, genetic analyses and oceanographic modelling of coastal and oceanic cod larval drift patterns were combined to elucidate the mechanisms responsible for an observed genetic cline over a >1500 km stretch along the coast of Norway. The results indicate that the north–south cline in coastal cod represents an extended contact zone between genetically divergent North Sea and Northeast Arctic cod and is maintained by two-way gene flow: by northward drift of pelagic eggs and larvae and by southward spawning migrations of Northeast Arctic cod. Computer simulations verify that the genetic cline can be established rapidly if gene flow into coastal populations is substantial. The shape of the cline, on the other hand, was found to be largely insensitive to the total amount of gene flow and therefore carries little information on extent of gene flow into and among coastal populations.

Many marine organisms have a permanent presence both inshore and offshore and spawn in multiple areas, yet their status as separate populations or stocks remain unclear. This is the situation for the northern shrimp (Pandalus borealis) around the Arctic Ocean, which in northern Norway represents an important income for a small-scale coastal fishery and a large-vessel offshore fleet. In Norwegian waters, we uncovered two distinct genetic clusters, viz. a Norwegian coastal and a Barents Sea cluster. Shrimps with a mixed heritage from the Norwegian coastal and the Barents Sea clusters, and genetically different from both, inhabit the fjords at the northernmost coast (Finnmark). Genetic structure between fjords did not display any general trend, and only the Varangerfjord in eastern Finnmark displayed significant genetic structure within the fjord. Shrimps in the Finnmark fjords differed in some degree from shrimps both in the adjacent Barents Sea and along the rest of the coast and should probably be considered a separate management unit.

Challenging long‐held perceptions of fish management units can help to protect vulnerable stocks. When a fishery consisting of multiple genetic stocks is managed as a single unit, overexploitation and depletion of minor genetic units can occur. Atlantic cod (Gadus morhua) is an economically and ecologically important marine species across the North Atlantic. The application of new genomic resources, including SNP arrays, allows us to detect and explore novel structure within specific cod management units. In Norwegian waters, coastal cod (i.e. those not undertaking extensive migrations) are divided into two arbitrary management units defined by ICES: one between 62° and 70°N (Norwegian coastal cod; NCC) and one between 58° and 62°N (Norwegian coastal south; NCS). Together, these capture a fishery area of >25,000 km2 containing many spawning grounds. To assess whether these geographic units correctly represent genetic stocks, we analysed spawning cod of NCC and NCS for more than 8,000 SNPs along with samples of Russian White Sea cod, north‐east Arctic cod (NEAC: the largest Atlantic stock), and outgroup samples representing the Irish and Faroe Sea's. Our analyses revealed large differences in spatial patterns of genetic differentiation across the genome and revealed a complex biological structure within NCC and NCS. Haplotype maps from four chromosome sets show regional specific SNP indicating a complex genetic structure. The current management plan dividing the coastal cod into only two management units does not accurately reflect the genetic units and needs to be revised. Coastal cod in Norway, while highly heterogenous, is also genetically distinct from neighbouring stocks in the north (NEAC), west (Faroe Island) and the south. The White Sea cod are highly divergent from other cod, possibly yielding support to the earlier notion of subspecies rank. publishedVersion

Identification of cryptic species can have profound implications in fishery management, conservation and biodiversity contexts. In the North Atlantic, the genus Sebastes is currently represented by four species, although additional cryptic species have been assumed. The connectivity of the gene-pools within the genus in Greenland waters, in particular, remains largely unexplored. Using a panel of 13 microsatellite markers for 720 fish, we explored the species complex of Sebastes norvegicus in Greenland waters. Genetic analyses provided evidence for three cryptic species in samples that were morphologically identified as S. norvegicus. They were termed S. norvegicus-A, S. norvegicus-B, and S. norvegicus-giants. A few phenotypic features exist to identify adult S. norvegicus giants, but no characteristics have been identified for the two other cryptic species. The proposed cryptic species should be recognized in the management regime to ensure sustainable exploitation and conservation of Sebastes species in Greenland waters.

Exploited marine resources can be managed more effectively when accurate information on geographic population structure is available. Genetic markers offer a powerful tool for fisheries management, because they reveal biologically sound management units. Single nucleotide polymorphisms (SNPs) markers derived from restriction-site associated DNA sequencing (RAD-seq) were developed and used to investigate the stock structure of Greenland halibut (Reinhardtius hippoglossoides). A total of 96 SNPs were analyzed from 384 individuals and eight locations across the Atlantic. Our results suggest a subdivision of Greenland halibut into two populations, an eastern Atlantic population and a western Atlantic population, with a proposed border across the Denmark Strait. In general, Greenland halibut display weak but significant population structure (overall FST= 0.003; p < 0.001), which can be explained by connectivity among populations owing to the migratory behavior or egg and larval drift.

The identification of isolated populations in widely distributed marine species is often impeded by low levels of genetic differentiation. However, modern genetic approaches now allow for the efficient detection of potentially large numbers of novel genetic variations, thereby improving the power to identify biologically meaningful management units. To investigate the genetic structuring of saithe (Pollachius virens L.), we applied seascape genetic approaches to 131 single nucleotide polymorphism loci genotyped in 584 fish collected from 11 locations across the North Atlantic. Wombling analysis and the Monmonier algorithm revealed four genetic clusters (Barents Sea, Central Northeast Atlantic, Rockall, and Canada) across the species distribution range. These results imply that genetic structuring in saithe may be influenced by abiotic factors such as geographical distance, and bathymetry as well as biotic factors such as sex-biased migration, and natal homing. The results suggest a potential mismatch between management and biological units across the Northeast Atlantic, which may have implications for sustainable exploitation of the species.

Genetic population structure is often used to identify management units in exploited species, but the extent of genetic differentiation may be inflated by geographic variation in the level of hybridization between species. We identify the genetic population structure of Sebastes mentella and investigate possible introgression within the genus by analyzing 13 microsatellites in 2562 redfish specimens sampled throughout the North Atlantic. The data support an historical divergence between the ‘shallow’ and ‘deep’ groups, beyond the Irminger Sea where they were described previously. A third group, ‘slope’, has an extended distribution on the East Greenland Shelf, in addition to earlier findings on the Icelandic Slope. Furthermore, S. mentella from the Northeast Arctic and Northwest Atlantic waters are genetically different populations. In both areas, interspecific introgression may influence allele frequency differences among populations. Evidence of introgression was found for almost all the identified Sebastes gene pools, but to a much lower extent than suggested earlier. Greenland waters appear to be a sympatric zone for many of the genetically independent Sebastes groups. This study illustrates that the identified groups maintain their genetic integrity in this region despite introgression. This article is protected by copyright. All rights reserved.

With the decline of many of the world’s fisheries, increased regulation, including marine protected areas (MPA), forms an increasingly important role in promoting sustainable resource use. Here, we present a novel “real-time” genetic monitoring programme used to protect the depleted Norwegian coastal cod stock (NCC) in an MPA during the spawning season, while a fishery targeted at the sustainable Northeast Arctic cod stock (NEAC) operates immediately outside. In the period 2009–2016, >6800 cod from the fishery were genotyped with the PanI locus that is discriminatory between these two stocks. The estimated fraction of NEAC increased during the study period until 2014; however, it did not exceed 70% for any sustained period. Therefore, the MPA remained closed for commercial harvest. Genetic analysis of eggs revealed a distinctly lower fraction of NEAC than in the catch from the adult stock, both immediately outside and within the MPA itself. We suggest that this discrepancy is likely to reflect differences in spawning areas used by NCC and NEAC. Estimated fractions of NEAC/NCC using PanI, otolith classification, and 39 single nucleotide polymorphisms were similar, thus validating the use of PanI to estimate NEAC/NCC composition.

The Northeast Arctic cod (Gadus morhua L.: NEAC) remains the most abundant cod stock in the North Atlantic, while the catches of the partially co-occurring Norwegian coastal cod (NCC) stocks have dramatically decreased in recent years. To ensure effective management of the two stocks, it is necessary to know if the population genetic structure is associated with any pattern in the spatial dynamics or whether it is affected by any distinct environmental factors. By combining information from electronic data storage tags (DST) and molecular genetics methods with statistical tools, we have been able to associate spatial dynamics and distinct environmental factors to the two cod stocks. In general, adult NEAC migrate between deep, warm overwintering grounds and shallow summer feeding grounds where water temperatures maybe low. In contrast, NCC do not undertake large-scale seasonal migrations, show little seasonal variation in depth distribution, and experience the opposite seasonal change in temperature compared with NEAC. However, within the NCC group, some individuals did conduct longer horizontal movements than others. Even though the distances calculated in this study represent the shortest distance between release and recapture positions, they are far higher than previously reported by NCC. Distinctive depth profiles indicate that this migrant NCC have moved out of the area, passing the deep trenches outside Lofoten while more stationary NCC occupies shallower depths throughout the year. The temperature profiles also indicate that migrant and stationary NCC has occupied different areas during the year. We demonstrate that the combination of information from DSTs and molecular genetics offers a deeper understanding of individual cod behaviour, provides an insight in the spatial dynamics of the species, and ultimately, improves the scientific basis for management of a complex mixed fishery of Atlantic cod.

We developed and characterized 20 microsatellite primer loci for the northern shrimp Pandalusborealis. All 20 loci were polymorphic with number of alleles ranging from 3 to 36 and with observed heterozygosity between 0.04 and 0.93. In addition, we tested the utility of these markers in three related shrimp species, P. montagui, Atlantopandaluspropinqvus and Dichelopandalusbonnieri. These new markers will prove useful in the identification of stock structure and hence, assessment of the commercially important species P. borealis.

The deepwater redfish Sebastes mentella has a wide distribution in the North Atlantic Ocean. In the Irminger Sea, there is evidence for two phenotypes (deep-sea and oceanic) of deepwater redfish. The two phenotypes have overlapping geographic distributions but differ in depth preferences. There are two hypotheses on deepwater redfish stock structure in the Irminger Sea. One suggests that mature individuals of a single stock segregate according to size and age and therefore that the phenotypes represent different life stages of the same stock. The other hypothesis suggests that there are two different stocks and that these stocks segregate mainly according to depth. Additionally, it is not clear whether the fish of the deep-sea phenotype in the Irminger Sea and those on the Icelandic continental shelf represent one stock. Analysis of genetic variability at eight allozyme markers in 1,763 deepwater redfish from 26 samples collected at different depths in the Irminger Sea and the Icelandic continent...

Hypoxia can influence fish growth, survival and on larger scales, population structure. These effects may be influenced by water temperature, and may vary intra-specifically with genotype. In Atlantic cod (Gadus morhua L.), the two haemoglobin homozygotes (Hb-I⁎11 and Hb-I⁎22) vary in oxygen affinity at different temperatures, which is thought to correspond to variation in hypoxia tolerance. We therefore tested if hypoxic avoidance behaviour in cod 1) depends on ambient temperature and 2) is modified by haemoglobin genotype. In a laminar flow choice box, we subjected juvenile cod to an initial phase of non-escapable hypoxia, and a subsequent recovery phase, where one habitat was kept at 20% O2 saturation while the other was raised in steps to full saturation. The experiment was performed at 5 and 15 °C with Hb-I⁎11 and Hb-I⁎22 cod. Cod responded to inescapable hypoxia by reducing their overall swimming speed and then, at the initial levels of the recovery phase, avoiding the most hypoxic habitat, irrespective of temperature or genotype. Fish recovered quickly as O2 levels increased, as evidenced by increased swimming speed and time spent in the most hypoxic habitat. The avoidance response depended strongly on temperature: the relative reduction in speed and avoidance of the most hypoxic habitat was more pronounced at 15 than at 5 °C. During the recovery phase, stressed fish initially maintained a higher swimming speed in the most hypoxic habitat. However, as O2 increased, swimming speed in both habitats converged. This point of convergence occurred at a lower O2 saturation at 5 °C. Fish ventilation rate in inescapable hypoxia was also higher at 15 °C. Haemoglobin genotype did not influence either ventilation rates or the nature of the hypoxic avoidance response at either temperature, but Hb-I⁎11 cod swam faster than Hb-I⁎22 cod in normoxia at 15 °C. Our results indicate that increased temperature limits the ability of cod of both haemoglobin genotypes to exploit hypoxic habitats. This may have negative future consequences for coastal cod stocks in light of increasing global temperatures and eutrophication in coastal waters. © 2008 Elsevier B.V. All rights reserved.

Atlantic cod is composed of multiple migratory and stationary populations widely distributed in the North Atlantic Ocean. The Northeast Arctic cod (NEAC) population in the Barents Sea undertakes annual spawning migrations to the northern Norwegian coast. Although spawning occurs sympatrically with the stationary Norwegian coastal cod (NCC), phenotypic and genetic differences between NEAC and NCC are maintained. In this study, we resolve the enigma by revealing the mechanisms underlying these differences. Extended linkage disequilibrium (LD) and population divergence were demonstrated in a 17.4-Mb region on linkage group 1 (LG1) based on genotypes of 494 SNPs from 192 parents of farmed families of NEAC, NCC or NEACxNCC crosses. Linkage analyses revealed two adjacent inversions within this region that repress meiotic recombination in NEACxNCC crosses. We identified a NEAC-specific haplotype consisting of 186 SNPs that was fixed in NEAC sampled from the Barents Sea, but segregating under Hardy–Weinberg equilibrium in eight NCC stocks. Comparative genomic analyses determine the NEAC configuration of the inversions to be the derived state and date it to ~1.6–2.0 Mya. The haplotype block harbours 763 genes, including candidates regulating swim bladder pressure, haem synthesis and skeletal muscle organization conferring adaptation to long-distance migrations and vertical movements down to large depths. Our results suggest that the migratory ecotype experiences strong directional selection for the two adjacent inversions on LG1. Despite interbreeding between NEAC and NCC, the inversions are maintaining genetic differentiation, and we hypothesize the co-occurrence of multiple adaptive alleles forming a ‘supergene’ in the NEAC population.

The large-scale population genetic structure of northern shrimp, Pandalus borealis, was investigated over the species’ range in the North Atlantic, identifying multiple genetically distinct groups. Genetic divergence among sample localities varied among 10 microsatellite loci (range: FST = −0.0002 to 0.0475) with a highly significant average (FST = 0.0149; P

The levels of heat-shock proteins of the 70 kDa family (Hsp70s) were measured in different soft tissues of Atlantic cod Gadus morhua from different locations and after exposure to various thermal conditions: acute temperature increments (1° C day -1 ), mid-term (73 days at 4-15° C) and long-term thermal acclimation (278 days at 8-15° C), and seasonal and latitudinal temperature variations (field samples). Tissue specific distribution patterns of Hsp70s were observed: liver > gills > red blood cells > brain > white muscle. Thus, different tissues may have required different levels of protection by Hsp70s, and possibly this was related to the rate of protein synthesis. There were no differences in tissue Hsp70s between Arctic cod populations (Arctic, i.e. Barents and White Seas, Norwegian coast, and North or Baltic Seas). No changes in Hsp70s levels were observed in response to temperature variation of any intensity (acute fluctuation or seasonal and latitudinal) within the range of physiological temperatures (4-15° C) in wild and laboratory Atlantic cod. This confirms previous observations that changes in Hsp70 caused by such temperature variation are often small in fishes. Probably, the constitutive level of Hsp70s in Atlantic cod was high enough to overcome potentially harmful effects of temperature variations within the physiological range. A suppressing effect of high temperature (15° C) has already been observed at a systematic level (as reduced rate of somatic growth), whereas it is not reflected in modified Hsp70s. Therefore, Hsp70s apparently played a secondary role in defining thermal tolerance limits in Atlantic cod. These conclusions are in line with a recent concept of thermal tolerance which indicated that the first line of thermal limitation in the cold and warm is a loss in aerobic scope.

Fourteen polymorphic microsatellite markers with tetranucleotide repeats were developed for the bearded goby (Sufflogobius bibarbatus) from partial genomic DNA libraries using a repeat enrichment protocol, and characterized using two putative populations from the northern Benguela. The average number of alleles per locus ranged from 4 to 34, and the observed heterozygosities across loci were between 0.237 and 0.983. We also tested the utility of these markers in two other marine gobies; sand goby Pomatoschistus minutus and two-spotted goby Gobiusculus flavescens. These polymorphic markers can be employed to investigate population structure and related questions of the bearded goby.

The genomic architecture underlying ecological divergence and ecological speciation with gene flow is still largely unknown for most organisms. One central question is whether divergence is genome-wide or localized in 'genomic mosaics' during early stages when gene flow is still pronounced. Empirical work has so far been limited, and the relative impacts of gene flow and natural selection on genomic patterns have not been fully explored. Here, we use ecotypes of Atlantic cod to investigate genomic patterns of diversity and population differentiation in a natural system characterized by high gene flow and large effective population sizes, properties which theoretically could restrict divergence in local genomic regions. We identify a genomic region of strong population differentiation, extending over approximately 20 cM, between pairs of migratory and stationary ecotypes examined at two different localities. Furthermore, the region is characterized by markedly reduced levels of genetic diversity in migratory ecotype samples. The results highlight the genomic region, or 'genomic island', as potentially associated with ecological divergence and suggest the involvement of a selective sweep. Finally, we also confirm earlier findings of localized genomic differentiation in three other linkage groups associated with divergence among eastern Atlantic populations. Thus, although the underlying mechanisms are still unknown, the results suggest that 'genomic mosaics' of differentiation may even be found under high levels of gene flow and that marine fishes may provide insightful model systems for studying and identifying initial targets of selection during ecological divergence.

Haemoglobins of 0-group cod from 30 to 150 mm length were analysed by agar gel electrophoresis and isoelectric focusing to identify the lower size limit for determination of the haemoglobin polymorphisms in cod; both the main system which has been known for more than 30 years and the recently described subgroups. The juvenile fish studied had higher concentration of the HbII-component (split into three bands at these fish sizes) compared to older fish. The main haemoglobin types could be distinguished clearly and reliably at a fish size of about 35 mm and above. The subgroups of the HbI-components, however, could not be identified reliably until the cod were about 80 mm.

To study the population structure of tusk Brosme brosme in the North Atlantic, samples from six fishing grounds were collected: St Kilda; Storegga; Faroe Islands; Rockall bank; Davis Strait and East Greenland. In addition two samples from the Norwegian coast; Kragero and Oygarden, were included. Nine enzymes and haemoglobin were analysed by starch gel and agar gel electrophoresis. In addition isoelectric focusing (IEF) was carried out. At least 22 loci were represented. Polymorphisms were shown in eight loci and average heterozygosity was found to be about 0.05. Highest heterozygosity was found in haemoglobin analysed by Smithies buffer pH 8.6. Malate dehydrogenase (MDH-2) analysed in IEF (pH 3.5-9.5), and haemoglobin were found as valuable tools for studies of stock structure in tusk. In the Greenland samples only one of the MDH-2 heterozygotes was found, while in the East Atlantic both the observed heterozygotes were more frequent. The frequencies of the the Hb*b allele were about 0.40 and 0.70 in the samples from east and west Atlantic respectively.

In a warmer Arctic, living conditions will change at all trophic levels of the marine ecosystem. Increased air and water temperatures will likely substantially reduce ice coverage. Trophic interactions might change and increased competition between resident Arctic species and invasive species seems likely. A theme session on “Marine harvesting in the Arctic” was held at the international Arctic Frontiers Conference in Tromsø, Norway, in January 2013. The theme session partitioned the topic into two sub-sessions: (i) introduced species, immigration and fate of resident species and (ii) prospective harvesting of marine biological resources in the Arctic. The four articles that follow this introduction are based on presentations made at the Arctic Frontiers theme session. These articles cover topics such as: how ice breeding seals (Pagophilus groenlandicus) can cope with ice retention in the Northwest Atlantic, how planktonic stages of the resident polar cod (Boreogadus saida) and the pole-ward expanding Pacific sand lance (Ammodytes hexapterus) may compete for food in the warming Beaufort Sea, and how the introduced red king crab (Paralithodes camtschaticus) disperse in the Barents Sea. The fourth article shows how differences in the life-history strategies of keystone zooplankton species will likely affect future productivity of commercial fisheries in polar regions.

14 páginas, 4 figuras.--Steve Cadrin ... et al.-- Acceso avanzado online., Throughout their range, Sebastes spp. are adapted to a diversity of ecological niches, with overlapping spatial distributions of different species that have little or no morphological differences. Divergence of behavioural groups into depth-defined adult habitats has led to reproductive isolation, adaptive radiation, and speciation in the genus Sebastes. Recent genetic research, supported by life-history information, indicates four biological stocks of Sebastes mentella in the Irminger Sea and adjacent waters: a western stock, a deep-pelagic stock, a shallow-pelagic stock, and an Iceland slope stock. Congruent differences in fatty acids and parasites suggest that these genetically distinct populations are adapted to disparate trophic habitats in pelagic waters (shallower and deeper than the deep-scattering layer) and in demersal habitats on the continental slope. Morphology of pelagic forms is also more streamlined than demersal forms. Although genetic differences and evidence for reproductive isolation are clear, these populations appear to share common nursery habitats on the Greenland shelf. We propose a redefinition of practical management units near the Irminger Sea based on geographic proxies for biological stocks and minimizing mixed-stock catches according to the spatial patterns of the recent fishery.

Population structuring in the North Atlantic deepwater demersal fish tusk (Brosme brosme) was studied with microsatellite DNA analyses. Screening eight samples from across the range of the species for seven loci revealed low but significant genetic heterogeneity (F(ST) = 0.0014). Spatial genetic variability was only weakly related to geographical (Euclidean) distance between study sites or separation of study sites along the path of major ocean currents. Instead, we found a significant effect of habitat, indicated by significant differentiation between relatively closely spaced sites: Rockall, which is surrounded by very deep water (1000 m), and the Mid-Atlantic Ridge, which is separated from the European slope by a deep ocean basin, were differentiated from relatively homogeneous sites across the Nordic Seas. Limited adult migration across bathymetric barriers in combination with limited intersite exchange of pelagic eggs and larvae due to site-specific circulatory retention or poor survival during drift phases across deep basins may be reducing gene flow. We regard these limitations to gene flow as the most likely mechanisms for the observed population structure in this demersal species. The results underscore the importance of habitat boundaries in marine species.

Background Marine fishes have been shown to display low levels of genetic structuring and associated high levels of gene flow, suggesting shallow evolutionary trajectories and, possibly, limited or lacking adaptive divergence among local populations. We investigated variation in 98 gene-associated single nucleotide polymorphisms (SNPs) for evidence of selection in local populations of Atlantic cod (Gadus morhua L.) across the species distribution. Results Our global genome scan analysis identified eight outlier gene loci with very high statistical support, likely to be subject to directional selection in local demes, or closely linked to loci under selection. Likewise, on a regional south/north transect of central and eastern Atlantic populations, seven loci displayed strongly elevated levels of genetic differentiation. Selection patterns among populations appeared to be relatively widespread and complex, i.e. outlier loci were generally not only associated with one of a few divergent local populations. Even on a limited geographical scale between the proximate North Sea and Baltic Sea populations four loci displayed evidence of adaptive evolution. Temporal genome scan analysis applied to DNA from archived otoliths from a Faeroese population demonstrated stability of the intra-population variation over 24 years. An exploratory landscape genetic analysis was used to elucidate potential effects of the most likely environmental factors responsible for the signatures of local adaptation. We found that genetic variation at several of the outlier loci was better correlated with temperature and/or salinity conditions at spawning grounds at spawning time than with geographic distance per se. Conclusion These findings illustrate that adaptive population divergence may indeed be prevalent despite seemingly high levels of gene flow, as found in most marine fishes. Thus, results have important implications for our understanding of the interplay of evolutionary forces in general, and for the conservation of marine biodiversity under rapidly increasing evolutionary pressure from climate and fisheries induced changes in local environments.

SUMMARYThe aim of this study was to determine the effects of seasonal temperature variation on the functional properties of lactate dehydrogenase (LDH) from white muscle and liver of Norwegian coastal cod (Gadus morhua) and the possible relevance of LDH allelic variability for thermal acclimation. Two groups of fishes were acclimated to 4°C or 12°C for one year. Polymorphism was observed in only one (Ldh-B) of the three Ldh loci expressed in cod liver and/or muscle. Isozyme expression remained unchanged regardless of acclimation temperature(TA). The products of locus Ldh-B comprise only 14–19% (depending on the tissue) of total LDH activities and,consequently, differences between phenotypes are negligible in terms of their effect on LDH total performance. No kinetic(\batchmode \documentclass[fleqn,10pt,legalpaper]{article} \usepackage{amssymb} \usepackage{amsfonts} \usepackage{amsmath} \pagestyle{empty} \begin{document} \(K_{\mathrm{m}}^{\mathrm{PYR}}\) \end{document}, \batchmode \documentclass[fleqn,10pt,legalpaper]{article} \usepackage{amssymb} \usepackage{amsfonts} \usepackage{amsmath} \pagestyle{empty} \begin{document} \(K_{\mathrm{si}}^{\mathrm{PYR}}\) \end{document}, Vmax) or thermodynamic (Ea,Δ G) differences were found among Ldh-B phenotypes. Clear kinetic differences were observed between LDH isoforms in the two tissues. However, the Arrhenius activation energy (Ea) for pyruvate reduction was the same for both tissues (Ea=47 kJ mol–1) at TA=12°C. Factors TA, tissue and phenotype did not reveal a significant effect on the Gibbs free energy change (ΔG) of the reaction(55.5 kJ mol–1). However, at TA=4°C,the Ea was increased (Ea=53–56 kJ mol–1) and the temperature dependence of the constant of substrate inhibition for pyruvate(\batchmode \documentclass[fleqn,10pt,legalpaper]{article} \usepackage{amssymb} \usepackage{amsfonts} \usepackage{amsmath} \pagestyle{empty} \begin{document} \(K_{\mathrm{si}}^{\mathrm{PYR}}\) \end{document}) decreased in both muscle and liver.In conclusion, the strategies of LDH adjustment to seasonal temperature variations in cod involve changes in LDH concentration (quantitative),adjustment of thermodynamic (Ea) and kinetic(\batchmode \documentclass[fleqn,10pt,legalpaper]{article} \usepackage{amssymb} \usepackage{amsfonts} \usepackage{amsmath} \pagestyle{empty} \begin{document} \(K_{\mathrm{si}}^{\mathrm{PYR}}\) \end{document}) properties of the LDH(modulative) but not the expression of alternative isoforms (qualitative). We assume that the observed increase in Ea and the decrease of temperature dependence of \batchmode \documentclass[fleqn,10pt,legalpaper]{article} \usepackage{amssymb} \usepackage{amsfonts} \usepackage{amsmath} \pagestyle{empty} \begin{document} \(K_{\mathrm{si}}^{\mathrm{PYR}}\) \end{document} at low TA is the result of structural changes of the LDH molecule(temperature-driven protein folding). We propose a new mechanism of metabolic compensation of seasonal temperature variations – cold acclimation results in changes in the kinetic and thermodynamic properties of LDH in a way that favours aerobic metabolism through reduction of the competition of LDH for pyruvate in normoxic conditions.

Activities of citrate synthase (CS), cytochrome c oxidase (CCO) and the electron transport system (ETS) were investigated in white muscle and liver of laboratory-maintained cod, Gadusmorhua, from the North Sea, Norwegian coast and Barents Sea for an analysis of temperature acclimation and adaptation in aerobic metabolism. Cold acclimation within each population led to elevated activities of CS, CCO and ETS in white muscle. In liver, however, only North Sea cod showed cold-compensated CS activities, with CCO and ETS unchanged. In contrast, cold-acclimated Norwegian cod displayed unchanged enzyme activities, and Arctic cod showed elevated activities for CS, but decreased activities for CCO and no change in ETS. Between-population comparisons revealed clear evidence for permanent cold adaptation in white muscle of northern (Norwegian coast and Barents Sea) compared to boreal (North Sea) cod populations, reflected by higher activities of CS, CCO and ETS at the same acclimation temperature. Cold-compensated, mass-specific enzyme activities in liver were found in northern compared to boreal cod for CS and CCO, however, for ETS, after warm acclimation only. When evaluated as capacity in total liver, such activities were only found in northern populations in the case of CS at all temperatures, and for CCO and ETS at 15°C only. Hepatosomatic index (IH) and liver lipid contents were highest in North Sea cod, with similar IH but lower lipid contents in cold- versus warm-acclimated animals. An acclimation effect on IH was found in Norwegian cod only, with higher IH but unchanged lipid contents in the cold. In conclusion, permanent cold adaptation of muscle aerobic metabolism prevails in cod populations at higher latitudes, which is in line with permanently elevated rates of oxygen consumption observed in a parallel study. These differences reflect higher maintenance costs in cold-adapted versus cold-acclimated cod.