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The artificial reefs in New York's waters provide structure in areas that are typically flat and sandy, creating habitat for a multitude of species as an area to spawn, forage, and reside. Passive acoustic data collected on the Fire Island and Shinnecock artificial reefs between 2018 and 2022 detected spawning-associated calls of weakfish (Cynoscion regalis) and Atlantic cod (Gadus morhua), as well as the presence of individual bottlenose dolphins (Tursiops truncatus) through their signature whistles. Weakfish and Atlantic cod were more vocally active on the Fire Island reef, where Atlantic cod grunts peaked during a new moon phase in December, and weakfish spawning experienced variable peaks between mid-July and mid-August on both reefs. Fifty-seven individual bottlenose dolphins were identified, with whistle repeats ranging from seconds to years apart. Passive acoustic monitoring allows for simultaneous collection of information on multiple species at different trophic levels as well as behavioral information that helps managers understand how these animals utilize these habitats, which can lead to improved conservation measures., (© 2024 Acoustical Society of America.)

Healthy ecosystems and species have some degree of resilience to changing conditions, however as the frequency and severity of environmental changes increase, resilience may be diminished or lost. In Sweden, one example of a species with reduced resilience is the Atlantic cod (Gadus morhua). This species has been subjected to overfishing, and with additional pressures such as habitat degradation and changing environmental conditions there has been little to no recovery, despite more than a decade of management actions. Given the historical ecological, economical, and cultural significance of cod, it is important to understand how Atlantic cod respond to global climate change to recover and sustainably manage this species in the future. A multi-stressor experiment was conducted to evaluate physiological responses of juvenile cod exposed to warming, ocean acidification, and freshening, changes expected to occur in their nursery habitat. The response to single drivers showed variable effects related to fish biometrics and increased levels of oxidative stress dependent parameters. Importantly, two separate responses were seen within a single treatment for the multi-stressor and freshening groups. These within-treatment differences were correlated to genotype, with the offshore ecotype having a heightened stress response compared to the coastal ecotype, which may be better adapted to tolerate future changes. These results demonstrate that, while Atlantic cod have some tolerance for future changes, ecotypes respond differently, and cumulative effects of multiple stressors may lead to deleterious effects for this important species., (© 2024. The Author(s).)

Increasing ocean temperature will speed up physiological rates of ectotherms. In fish, this is suggested to cause earlier spawning due to faster oocyte growth rates. Over time, this could cause spawning time to become decoupled from the timing of offspring food resources, a phenomenon referred to as trophic asynchrony. We used biological data, including body length, age, and gonad developmental stages collected from >125,000 individual Northeast Arctic cod (Gadus morhua) sampled between 59 and 73° N in 1980-2019. Combined with experimental data on oocyte growth rates, our analyses show that cod spawned progressively earlier by about a week per decade, partly due to ocean warming. It also appears that spawning time varied by more than 40 days, depending on year and spawning location. The significant plasticity in spawning time seems to be fine-tuned to the local phytoplankton spring bloom phenology. This ability to partly overcome thermal drivers and thus modulate spawning time could allow individuals to maximize fitness by closely tracking local environmental conditions important for offspring survival. Our finding highlights a new dimension for trophic match-mismatch and should be an important consideration in models used to predict phenology dynamics in a warmer climate., (© 2024 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Ecological resilience is the capability of an ecosystem to maintain the same structure and function and avoid crossing catastrophic tipping points (i.e. undergoing irreversible regime shifts). While fundamental for management, concrete ways to estimate and interpret resilience in real ecosystems are still lacking. Here, we develop an empirical approach to estimate resilience based on the stochastic cusp model derived from catastrophe theory. The cusp model models tipping points derived from a cusp bifurcation. We extend cusp in order to identify the presence of stable and unstable states in complex natural systems. Our Cusp Resilience Assessment (CUSPRA) has three characteristics: (i) it provides estimates on how likely a system is to cross a tipping point (in the form of a cusp bifurcation) characterized by hysteresis, (ii) it assesses resilience in relation to multiple external drivers and (iii) it produces straightforward results for ecosystem-based management. We validate our approach using simulated data and demonstrate its application using empirical time series of an Atlantic cod population and marine ecosystems in the North Sea and the Mediterranean Sea. We show that Cusp Resilience Assessment is a powerful method to empirically estimate resilience in support of a sustainable management of our constantly adapting ecosystems under global climate change.

At high latitudes, the suitable window for timing reproductive events is particularly narrow, promoting tight synchrony between trophic levels. Climate change may disrupt this synchrony due to diverging responses to temperature between, for example, the early life stages of higher trophic levels and their food resources. Evidence for this is equivocal, and the role of compensatory mechanisms is poorly understood. Here, we show how a combination of ocean warming and coastal water darkening drive long-term changes in phytoplankton spring bloom timing in Lofoten Norway, and how spawning time of Northeast Arctic cod responds in synchrony. Spring bloom timing was derived from hydrographical observations dating back to 1936, while cod spawning time was estimated from weekly fisheries catch and roe landing data since 1877. Our results suggest that land use change and freshwater run-off causing coastal water darkening has gradually delayed the spring bloom up to the late 1980s after which ocean warming has caused it to advance. The cod appear to track phytoplankton dynamics by timing gonadal development and spawning to maximize overlap between offspring hatch date and predicted resource availability. This finding emphasises the importance of land-ocean coupling for coastal ecosystem functioning, and the potential for fish to adapt through phenotypic plasticity., (© 2024 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Marine vibrators are a new technology being developed for seismic surveys. These devices can transmit continuous instead of impulsive sound and operate over a narrower frequency band and at lower peak pressure than airguns, which is assumed to reduce their environmental impacts. We exposed spawning Atlantic cod (Gadus morhua) to sound produced by a prototype, but full-scale, marine vibrator, and monitored behavioural responses of tagged cod using acoustic telemetry. Fish were exposed to 10 × 3 h continuous sound treatments over a 4-day period using a randomised-block design. Sound exposure levels were comparable to airgun exposure experiments conducted previously with the same set-up ranging from ∼115 to 145 dB re 1 μPa 2 s during exposure. Telemetry data were used to assess 1) whether marine vibrator exposure displaced cod from the spawning ground, through estimation of residence and survival probabilities, and 2) fine-scale behavioural responses within the test site, namely swimming depth, activity levels, displacement, and home ranges. Forty-two spawning cod were tagged prior to the exposure, with 22 present during the exposure. All 22 tags were equipped with pressure sensors and ten of these additionally with accelerometers. While no premature departure from the spawning site was observed, cod reacted to the exposure by decreasing their activity levels (by up to 50%, SE = 7%) and increasing their swimming depth (by up to 2.5 m, SE = 1.0 m) within the test site during the exposure period. These behavioural responses varied by sex and time of day. Cod reactions to a marine vibrator may be more pronounced than reactions to airguns, possibly because continuous sound is more disturbing to fish than intermittent sound at the same exposure levels. However, given sample size limitations of the present study, further studies with continuous sound are necessary to fully understand its impact and biological significance., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Microplastics (MPs) have become a global issue as they are omnipresent in the ocean. Fish ingesting MPs through feed could be affected in their physiological function, e.g., disrupted enzyme production and function, reduction of feeding and reproductive failure. This study assessed the effects of feed containing naturally weathered MPs from the Oslofjord (Norway) on the reproductive physiology of Atlantic cod (Gadus morhua). Farmed cod broodstock were fed either control (C-diet) or feeds containing 1% microplastic (MP-diet) starting nine months prior to spawning, from June until May. No major differences were found between diet groups in overall biometrics or gonad histology. Sex steroid levels (testosterone, 11-ketotestosterone and 17β-estradiol) resulted in expected profiles increasing over time without any significant differences between treatments. Gene expression levels of the steroidogenic enzyme 20β-hydroxysteroid dehydrogenase (20β-hsd) and vitellogenin1 (vtg1) showed significant differences between dietary treatments with lower expression in the control group. This can be a direct effect of MPs, but endocrine disrupting effects of potentially leachable plastic additives cannot be completely ruled out. Thus, these enzymes could be indicators of exposure to contaminants that disrupt sexual maturation by affecting the production of primarily maturation-inducing steroid. Although the concentration of MPs employed in this study may not be high enough to elicit any observable short-term biological effects, the observed gene expression suggests that long-term consequences should be considered caused by an expected increase of MPs in marine environments., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Long-term changes in the age and size structure of animal populations are well documented, yet their impacts on population productivity are poorly understood. Fishery exploitation can be a major driver of changes in population age-size structure because fisheries significantly increase mortality and often selectively remove larger and older fish. Climate change is another potential driver of shifts in the demographic structure of fish populations. Northeast Arctic (NEA) cod is the largest population of Atlantic cod (Gadus morhua) and one of the world's most important commercial fish stocks. This population has experienced considerable changes in population age-size structure over the past century, largely in response to fishing. In this study, we investigate whether changes in spawner age structure have affected population productivity in NEA cod, measured as recruits per spawning stock biomass, over the past 75 years. We find evidence that shifts in age structure toward younger spawners negatively affect population productivity, implying higher recruitment success when the spawning stock is composed of older individuals. The positive effect of an older spawning stock is likely linked to maternal effects and higher reproductive output of larger females. Our results indicate a threefold difference in productivity between the youngest and oldest spawning stock that has been observed since the 1950s. Further, our results suggest a positive effect of environmental temperature and a negative effect of intraspecific cannibalism by older juveniles on population productivity, which partly masked the effect of spawner age structure unless accounted for in the model. Collectively, these findings emphasize the importance of population age structure for the productivity of fish populations and suggest that harvest-induced demographic changes can have negative feedbacks for fisheries that lead to a younger spawning stock. Incorporating demographic data into harvest strategies could thus facilitate sustainable fishery management., (© 2022 The Ecological Society of America.)

Sustainable human exploitation of living marine resources stems from a delicate balance between yield stability and population persistence to achieve socioeconomic and conservation goals. But our imperfect knowledge of how oceanic oscillations regulate temporal variation in an exploited species can obscure the risk of missing management targets. We illustrate how applying a management policy to suppress fluctuations in fishery yield in variable environments (prey density and regional climate) can present unintended outcomes in harvested predators and the sustainability of harvesting. Using Atlantic cod (Gadus morhua, an apex predatory fish) in the Barents Sea as a case study we simulate age-structured population and harvest dynamics through time-varying, density-dependent and density-independent processes with a stochastic, process-based model informed by 27-year monitoring data. In this model, capelin (Mallotus villosus, a pelagic forage fish), a primary prey of cod, fluctuations modulate the strength of density-dependent regulation primarily through cannibalistic pressure on juvenile cod survival; sea temperature fluctuations modulate thermal regulation of cod feeding, growth, maturation, and reproduction. We first explore how capelin and temperature fluctuations filtered through cod intrinsic dynamics modify catch stability and then evaluate how management to suppress short-term variability in catch targets alters overharvest risk. Analyses revealed that suppressing year-to-year catch variability impedes management responses to adjust fishing pressure, which becomes progressively out of sync with variations in cod abundance. This asynchrony becomes amplified in fluctuating environments, magnifying the amplitudes of both fishing pressure and cod abundance and then intensifying the density-dependent regulation of juvenile survival through cannibalism. Although these transient dynamics theoretically give higher average catches, emergent, quasicyclic behaviors of the population would increase long-term yield variability and elevate overharvest risk. Management strategies that overlook the interplay of extrinsic (fishing and environment) and intrinsic (life history and demography) fluctuations thus can inadvertently destabilize fish stocks, thereby jeopardizing the sustainability of harvesting. These policy implications underscore the value of ecosystem approaches to designing management measures to sustainably harvest ecologically connected resources while achieving socioeconomic security., (© 2021 The Authors. Ecological Applications published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

According to the theory of compensatory dynamics, depleted populations should recover when the threat responsible for their decline is removed because per capita population growth is assumed to be highest when populations are at their smallest viable sizes. Yet, many seriously depleted fish populations have failed to recover despite threat mitigation. Atlantic cod ( Gadus morhua ) stocks off Newfoundland, despite 30 years of dramatically reduced fishing mortality and numerous fishery closures, have not recovered, suggesting that drivers other than fishing can regulate the growth of collapsed fish populations, inhibiting or preventing their recovery. Here, using Bayesian inference, we show strong evidence of Allee effects in a south Newfoundland cod population, based on data on recruitment and spawning stock biomass. We infer the Allee-effect threshold, below which recovery is impaired. We demonstrate the necessity of data at low population sizes to make inferences about the nature of low-abundance dynamics. Our work indicates that Allee effects are not negligible in commercially exploited fish populations, as commonly projected, and that they represent an inhibitory force that can effectively prevent recovery from overfishing. Our findings contrast with prevailing fisheries management practices that assume compensatory dynamics at low abundances with potential to seriously overestimate the recovery potential of collapsed populations.

Evidence from mammals and aves alludes to a possibly conserved seasonal photoperiod induced neuroendocrine cascade which stimulates subsequent sexual maturation however our understanding of this mechanism in teleosts is lacking. Unlike all teleosts studied to date, the Atlantic cod (Gadus morhua) is a short day breeder with the reduction in day-length from the summer solstice stimulating gametogenesis. Cod specific orthologues of eya3, tshβ and dio2 were identified and their expression was monitored in the brain and pituitary of cod held under either stimulated or inhibited photoperiod conditions. While no differential expression was apparent in brain dio2 & tshβ and pituitary tshβ, there was significant temporal variation in expression of pituitary eya3 under the SNP treatment, with expression level elevating in association with active gametogenesis. Under the LL treatment, sexual maturation was inhibited and there was a corresponding suppression of eya3 expression. In a second study the impact of size/energetic status on the initiation of sexual maturation was investigated. In the feed restricted population maturation was significantly suppressed (5% sexually mature) compared to the ab libitum fed stock (95% sexually mature) with there being a concomitant significant suppression in pituitary eya3 expression. Overall, these results suggest that pituitary eya3 has the potential to act as an integrator of both environmental and energetic regulation of sexual maturation of cod. Being the first account of eya3 induction in a short day breeding teleost, the conserved association with stimulation of reproduction and not seasonal state indicates that the upstream drivers which initiate the pathway differ among vertebrates according to their breeding strategies, but the pathway itself and its role in the reproductive cascade appears to be conserved across the vertebrate clade., (Copyright © 2021 Elsevier Inc. All rights reserved.)

Specific changes identified in the otolith macrostructure of Northeast Arctic cod as "spawning zones" are presumed to represent spawning events, but recent experimental studies have challenged this relationship. Because these zones are not routinely recorded outside of Norway, otoliths from multiple Atlantic cod populations with different life history and environmental traits were first examined to see if spawning zones could be identified as a general characteristic of cod. Then, a large archival collection of cod otoliths was used to investigate temporal changes in the occurrence of spawning zones and test for correlations between maturity at age derived from otolith spawning zones and gonad maturity stages. This study shows that spawning zones likely are a universal trait of Atlantic cod and not limited to certain environments or migratory behaviors as previously proposed. Maturity at age derived from spawning zone data showed trends consistent with those from gonad examinations. However, spawning zones appear to form with a one- or two-year lag with sexual maturity, which is suspected to reflect a stabilizing of energy partitioning after the first spawning events. Our results illustrate the potential for use of spawning zones, for example in species or populations with limited available maturity data, and highlights the need for addressing the physiological processes behind their formation., Competing Interests: The authors have declared that no competing interests exist.

The mode and extent of rapid evolution and genomic change in response to human harvesting are key conservation issues. Although experiments and models have shown a high potential for both genetic and phenotypic change in response to fishing, empirical examples of genetic responses in wild populations are rare. Here, we compare whole-genome sequence data of Atlantic cod ( Gadus morhua ) that were collected before (early 20th century) and after (early 21st century) periods of intensive exploitation and rapid decline in the age of maturation from two geographically distinct populations in Newfoundland, Canada, and the northeast Arctic, Norway. Our temporal, genome-wide analyses of 346,290 loci show no substantial loss of genetic diversity and high effective population sizes. Moreover, we do not find distinct signals of strong selective sweeps anywhere in the genome, although we cannot rule out the possibility of highly polygenic evolution. Our observations suggest that phenotypic change in these populations is not constrained by irreversible loss of genomic variation and thus imply that former traits could be reestablished with demographic recovery., Competing Interests: The authors declare no competing interest., (Copyright © 2021 the Author(s). Published by PNAS.)

Studying biology of sperm provides valuable information to optimize artificial reproduction and is crucial for sustainable aquaculture. Here, we investigated morphology of spermatozoon in Atlantic cod (Gadus morhua) using transmission and scanning electron microscopy. Furthermore, spermatozoa motility kinetics at different osmolalities were studied using computer-assisted sperm analysis software. The spermatozoon lacked an acrosome and consisted of a head, midpiece, and flagellum. The head of spermatozoa was round, oval, and rather elongated in shape, showing high variations in dimensions. There were up to 6 mitochondria that encircled the proximal part of the flagellum. The proximal and distal centrioles were located within the nuclear notch and arranged orthogonal to each other. The axoneme had a typical 9 + 2 microtubule structure. The flagellar length of spermatozoon was 66.94 ± 0.46 μm. Spermatozoa were immotile in the seminal plasma. Dilution of sperm with natural seawater (1100 mOsmol/kg) resulted in initiation of motility for 91.0 ± 3.4% of spermatozoa with average velocity of 86.2 ± 2.3 μm/s and beating frequency of 52 Hz. The duration of spermatozoa motility was > 6 min; however, the percentage of motile spermatozoa decreased at 60 s post-activation. When osmolality of natural seawater was modified using distilled water or NaCl, spermatozoa motility was not initiated at ≤ 400 and ≥ 2500 mOsmol/kg, and the highest percentage of motility was observed at 730-1580 mOsmol/kg. In a sucrose solution, spermatozoa motility was initiated and suppressed at 600 and 1500 mOsmol/kg, respectively, and highest percentage of motility was observed at 800-1100 mOsmol/kg. Spermatozoon morphology comparisons within Gadiformes showed differences in dimensions of head and mitochondria, flagellar length, and number of mitochondria. The present study provides valuable data that can be used for phylogenetic implications based on spermatozoon morphology and for development of artificial fertilization and sperm cryopreservation protocols based on sperm motility.

This study presents the diet composition of western Baltic cod Gadus morhua based on 3150 stomachs sampled year-round between 2016 and 2017 using angling, gillnetting and bottom trawling, which enhanced the spatio-temporal coverage of cod habitats. Cod diet composition in shallow areas (<20 m depth) was dominated by benthic invertebrate species, mainly the common shore crab Carcinus maneas. Compared to historic diet data from the 1960s and 1980s (limited to depth >20 m), the contribution of herring Clupea harengus decreased and round goby Neogobius melanostomus occurred as a new prey species. Statistical modelling revealed significant relationships between diet composition, catch depth, fish length and season. Generalized additive modelling identified a negative relationship between catch depth and stomach content weight, suggesting reduced food intake in winter when cod use deeper areas for spawning and during peak summer when cod tend to avoid high water temperatures. The results of this study highlight the importance of shallow coastal areas as major feeding habitats of adult cod in the western Baltic Sea, which were previously unknown because samples were restricted to deeper trawlable areas. The results strongly suggest that historic stomach analyses overestimated the role of forage fish and underestimated the role of invertebrate prey. Eventually, this study shows the importance of a comprehensive habitat coverage for unbiased stomach sampling programmes to provide a more reliable estimation of top predator diet, a key information for food web analyses and multispecies models., (© 2020 The Authors. Journal of Fish Biology published by John Wiley & Sons Ltd on behalf of The Fisheries Society of the British Isles.)

Seasonal prey bursts are important for the life cycles and energy budgets of many predators. This study documents the diet and, especially, the importance of the ephemeral occurrence of capelin as prey for Atlantic cod (Gadus morhua) in Godthaabsfjord, west Greenland, over an annual cycle. The cod showed clear differences in diet composition on the 11 sampling dates resulting in a spring-summer, late summer-autumn and winter cluster. Moreover, a single sampling date, 12 May, was defined by cod gorge feeding on spawning capelin, which led to average stomach contents 4.3 times higher than the average for the remaining sampling dates. Changes in nitrogen stable isotope values from 22 April to 7 July in cod liver and muscle tissue were used to calculate the consumption of capelin. Based on this, the consumption of capelin varied between 538 and 658 g wet weight for a 1.3 kg cod. Using published consumption/biomass estimates and observed growth rates, the capelin intake corresponds to 10.1%-33.3% of the annual food consumption and accounts for 28.1%-34.5% of the annual growth of the cod. The present study documents the omnivorous feeding mode of Atlantic cod but highlights the utilization and importance of ephemeral prey bursts for the annual energy budget of the cod. It is hypothesized that access to capelin is critical for the postspawning recovery of Godthaabsfjord cod., (© 2020 Fisheries Society of the British Isles.)

Fish otoliths' chronometric properties make them useful for age and growth rate estimation in fisheries management. For the Eastern Baltic Sea cod stock (Gadus morhua), unclear seasonal growth zones in otoliths have resulted in unreliable age and growth information. Here, a new age estimation method based on seasonal patterns in trace elemental otolith incorporation was tested for the first time and compared with the traditional method of visually counting growth zones, using otoliths from the Baltic and North seas. Various trace elemental ratios, linked to fish metabolic activity (higher in summer) or external environment (migration to colder, deeper habitats with higher salinity in winter), were tested for age estimation based on assessing their seasonal variations in concentration. Mg:Ca and P:Ca, both proxies for growth and metabolic activity, showed greatest seasonality and therefore have the best potential to be used as chemical clocks. Otolith image readability was significantly lower in the Baltic than in the North Sea. The chemical (novel) method had an overall greater precision and percentage agreement among readers (11.2%, 74.0%) than the visual (traditional) method (23.1%, 51.0%). Visual readers generally selected more highly contrasting zones as annuli whereas the chemical readers identified brighter regions within the first two annuli and darker zones thereafter. Visual estimates produced significantly higher, more variable ages than did the chemical ones. Based on the analyses in our study, we suggest that otolith microchemistry is a promising alternative ageing method for fish populations difficult to age, such as the Eastern Baltic cod., (© 2020 The Authors. Journal of Fish Biology published by John Wiley & Sons Ltd on behalf of The Fisheries Society of the British Isles.)

Anthropogenic underwater noise may negatively affect marine animals. Yet, while fishes are highly sensitive to sounds, effects of acoustic disturbances on fishes have not been extensively studied at the population level. In this study, we use a size-structured model based on energy budgets to analyse potential population-level effects of anthropogenic noise on Atlantic cod ( Gadus morhua ). Using the model framework, we assess the impact of four possible effect pathways of disturbance on the cod population growth rate. Through increased stress, changes in foraging and movement behaviour, and effects on the auditory system, anthropogenic noise can lead to (i) increased energy expenditure, (ii) reduced food intake, (iii) increased mortality, and (iv) reduced reproductive output. Our results show that population growth rates are particularly sensitive to changes in energy expenditure and food intake because they indirectly affect the age of maturation, survival and fecundity. Sub-lethal effects of sound exposure may thus affect populations of cod and fishes with similar life histories more than lethal effects of sound exposure. Moreover, anthropogenic noise may negatively affect populations when causing persistent increases of energy expenditure or decreases of food intake. Effects of specific acoustic pollutants on energy acquisition and expenditure should therefore be further investigated.

The collection and presentation of accurate reproductive data from wild fish has historically been somewhat problematic, especially for serially spawning species. Therefore, the aim of the current study was to develop a novel method of assessing female spawning status that is robust to variation in oocyte dynamics between specimens. Atlantic cod (Barents Sea stock) were used to develop the new 'ultrametric' method, that is based on the progressive depletion of the vitellogenic oocyte pool relative to the rather constant previtellogenic oocyte (PVO) pool. Fish were subsequently partitioned into one of four categories that accurately reflected changes in their oocyte size frequency distribution characteristics and gonadosomatic index throughout spawning. The ultrametric method overcomes difficulties associated with presence of bimodal oocyte distributions, oocyte tails, lack of clear hiatus region, and presence of free ova, and can be implemented at a single sampling point. Much of the workflow is fully automated, and the technique may circumvent the need for histological analysis depending on the desired outcome. The ultrametric method differs from the traditional autodiametric method in that PVOs can be separated by ultrasonication and then enumerated, and ovarian homogeneity is not a mandatory requirement per se. The method is designed for determinate spawners but might be extended to include indeterminate spawners.

The Arctic Ocean is an early warning system for indicators and effects of climate change. We use a novel combination of experimental and time-series data on effects of ocean warming and acidification on the commercially important Northeast Arctic cod (Gadus morhua) to incorporate these physiological processes into the recruitment model of the fish population. By running an ecological-economic optimization model, we investigate how the interaction of ocean warming, acidification and fishing pressure affects the sustainability of the fishery in terms of ecological, economic, social and consumer-related indicators, ranging from present day conditions up to future climate change scenarios. We find that near-term climate change will benefit the fishery, but under likely future warming and acidification this large fishery is at risk of collapse by the end of the century, even with the best adaptation effort in terms of reduced fishing pressure., Competing Interests: The authors have declared that no competing interests exist.

Physiological mechanisms determining thermal limits in fishes are debated but remain elusive. It has been hypothesised that motor function loss, observed as loss of equilibrium during acute warming, is due to direct thermal effects on brain neuronal function. To test this, we mounted cooling plates on the heads of Atlantic cod ( Gadus morhua ) and quantified whether local brain cooling increased whole-organism acute upper thermal tolerance. Brain cooling reduced brain temperature by 2-6°C below ambient water temperature and increased thermal tolerance by 0.5 and 0.6°C on average relative to instrumented and uninstrumented controls, respectively, suggesting that direct thermal effects on brain neurons may contribute to setting upper thermal limits in fish. However, the improvement in thermal tolerance with brain cooling was small relative to the difference in brain temperature, demonstrating that other mechanisms (e.g. failure of spinal and peripheral neurons, or muscle) may also contribute to controlling acute thermal tolerance., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

While many in-laboratory ecotoxicological studies have shown the adverse impact of pollutants to the fitness of an individual, direct evidence from the field on the population dynamics of wildlife animals has been lacking. Here, we provide empirical support for a negative effect of pollution on Atlantic cod (Gadus morhua) population dynamics in coastal waters of Norway by combining unique time series of juvenile cod abundance, body size, environmental concentration of toxic contaminants and a spatially structured population dynamics model. The study shows that mercury concentration might have decreased the reproductive potential of cod in the region despite the general decline in the environmental concentration of mercury, cadmium and hexachlorobenzene since the implementation of national environmental laws. However, some cod populations appeared to be more resistant to mercury pollution than others, and the strength and shape of mercury effect on cod reproductive potential was fjord-specific. Additionally, cod growth rate changed at scales smaller than fjords with a gradient related to the exposure to the open ocean and offshore cod. These spatial differences in life-history traits emphasize the importance of local adaptation in shaping the dynamics of local wildlife populations. Finally, this study highlights the possibility to mitigate pollution effects on natural populations by reducing the overall pollution level, but also reveals that pollution reduction alone is not enough to rebuild local cod populations. Cod population recovery probably requires complementary efforts on fishing regulation and habitat restoration.

A multitude of recent studies have documented the detrimental effects of crude oil exposure on early life stages of fish, including larvae and embryos. While polycyclic aromatic hydrocarbons (PAHs), particularly alkyl PAHs, are often considered the main cause of observed toxic effects, other crude oil derived organic compounds are usually overlooked. In the current study, comprehensive two-dimensional gas chromatography coupled to mass spectrometry was applied to investigate the body burden of a wide range of petrogenic compounds in Atlantic haddock (Melanogrammus aeglefinus) and cod (Gadus morhua) embryos that had been exposed to sublethal doses of dispersed crude oil. Several groups of alkylated monoaromatic compounds (e.g. alkyl tetralins, indanes and alkyl benzenes), as well as highly alkylated PAHs, were found to accumulate in the fish embryos upon crude oil exposure. To investigate the toxicity of the monoaromatic compounds, two models (1-isopropyl-4-methyltetralin and 1-isopropyl-4-methylindane) were synthesized and shown to bioaccumulate and cause delayed hatching in developing embryos. Minor developmental effects, including craniofacial and jaw deformations and pericardial edemas, were also observed at the highest studied concentrations of the alkylindane., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Atlantic cod is a species that is affected by climate change, with some populations being exposed to higher temperatures than others. The two polymorphs of its major haemoglobin type (HbI) show an inverse change in frequency along a latitudinal temperature cline in the North East Atlantic, which has been associated with differences in population performance and behavioural traits. An earlier study at the northern distribution limit of the species reported differential temperature sensitivities of red blood cell oxygen (O 2 ) affinity between the northern cold-water HbI-2 polymorph and its southern, warm-water HbI-1 counter-part, which has since widely been held as adaptive for the species across its distributional range. The present study critically re-examined this hypothesis by comparing the thermal sensitivity of O 2 binding in both purified HbI polymorphs from the southern, high-temperature distribution limit of the species under controlled conditions of allosteric modifiers of Hb function. Contrary to the prevailing view, the O 2 affinity of the major HbI polymorphs did not differ from each other under any of the tested conditions. Depending on pH and ATP concentration, the temperature-sensitive and temperature-insensitive Hb-O 2 affinity phenotypes - previously exclusively ascribed to HbI-1 and HbI-2, respectively - could be induced in both HbI polymorphs. These results are the first to establish a molecular mechanism behind a reversed temperature dependence of red blood cell O 2 affinity in a non-endotherm fish and lay the basis for future studies on alternative mechanisms behind the differences in distribution, performance and behavioural traits associated with the different HbI polymorphs of Atlantic cod., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

Exploitation and changing ocean conditions have resulted in altered species interactions and varied population dynamics within marine fish communities off northeast Newfoundland and southern Labrador, Canada. To understand contemporary species interactions, we quantified the isotopic niches, niche overlap, and ontogenetic niche change among seven dominant fish species using stable isotope analyses. Analyses used fishes from three regions differing in fish and prey diversities. Differences in fish and diet composition diversity among regions were found using Simpson's inverse diversity index. The regions of lowest diversities had higher instances of niche overlap and higher percentage of niche overlap area. The region of highest diversity had the widest spread of niches with greater distances from the community centroid. Ontogenetic shifts were observed such that larger individuals shifted towards the community centroid with the exception of Atlantic cod. Atlantic cod in particular was found to consistently be the top predator of the analyzed species. Our results reveal: (a) overlap in isotopic niches and spread within niche space was correlated with fish and diet diversity; (b) ontogenetic shifts are important when considering a species' niche and quantifying spatial variation in community niche profiles., Competing Interests: The authors have declared that no competing interests exist.

Introgressive hybridization creates networks of genetic relationships across species. Among marine fish of the Gadidae family, Pacific cod and walleye pollock are separate invasions of an Atlantic cod ancestor into the Pacific. Cods are ecological success stories, and their ecologies allow them to support the largest fisheries of the world. The enigmatic walleye pollock differs morphologically, behaviorally, and ecologically from its relatives, representing a niche shift. Here, we apply whole-genome sequencing to Pacific, Arctic, and Atlantic gadids and reveal extensive introgression among them with the ABBA-BABA test and pseudolikelihood phylogenetic network analysis. We propose that walleye pollock resulted from extensive adaptive introgression or homoploid hybrid speciation. The path of evolution of these taxa is more web than a tree. Their ability to invade and expand into new habitats and become ecologically successful may depend on genes acquired through adaptive introgression or hybrid speciation.

Collapses and regime changes are pervasive in complex systems (such as marine ecosystems) governed by multiple stressors. The demise of Atlantic cod ( Gadus morhua) stocks constitutes a text book example of the consequences of overexploiting marine living resources, yet the drivers of these nearly synchronous collapses are still debated. Moreover, it is still unclear why rebuilding of collapsed fish stocks such as cod is often slow or absent. Here, we apply the stochastic cusp model, based on catastrophe theory, and show that collapse and recovery of cod stocks are potentially driven by the specific interaction between exploitation pressure and environmental drivers. Our statistical modelling study demonstrates that for most of the cod stocks, ocean warming could induce a nonlinear discontinuous relationship between fishing pressure and stock size, which would explain hysteresis in their response to reduced exploitation pressure. Our study suggests further that a continuing increase in ocean temperatures will probably limit productivity and hence future fishing opportunities for most cod stocks of the Atlantic Ocean. Moreover, our study contributes to the ongoing discussion on the importance of climate and fishing effects on commercially exploited fish stocks, highlighting the importance of considering discontinuous dynamics in holistic ecosystem-based management approaches, particularly under climate change.

In order to understand the effect of global change on marine fishes, it is imperative to quantify the effects on fundamental parameters such as survival and growth. Larval survival and recruitment of the Atlantic cod (Gadus morhua) were found to be heavily impaired by end-of-century levels of ocean acidification. Here, we analysed larval growth among 35-36 days old surviving larvae, along with organ development and ossification of the skeleton. We combined CO 2 treatments (ambient: 503 µatm, elevated: 1,179 µatm) with food availability in order to evaluate the effect of energy limitation in addition to the ocean acidification stressor. As expected, larval size (as a proxy for growth) and skeletogenesis were positively affected by high food availability. We found significant interactions between acidification and food availability. Larvae fed ad libitum showed little difference in growth and skeletogenesis due to the CO 2 treatment. Larvae under energy limitation were significantly larger and had further developed skeletal structures in the elevated CO 2 treatment compared to the ambient CO 2 treatment. However, the elevated CO 2 group revealed impairments in critically important organs, such as the liver, and had comparatively smaller functional gills indicating a mismatch between size and function. It is therefore likely that individual larvae that had survived acidification treatments will suffer from impairments later during ontogeny. Our study highlights important allocation trade-off between growth and organ development, which is critically important to interpret acidification effects on early life stages of fish., (© 2018 John Wiley & Sons Ltd.)

The aim of this study was to assess whether fish in Kollevåg, a sheltered bay on the western coast of Norway, previously utilized as a waste disposal site, could be affected by environmental contaminants leaking from the waste. Farmed, juvenile Atlantic cod (Gadus morhua) were caged for six weeks at three different locations in Kollevåg bay and at one reference location. Sediments and cod samples (bile and liver) were analyzed for polychlorinated biphenyls (PCBs), organochlorine pesticides (OCPs), brominated flame retardants (BFRs), per-and polyfluoroalkyl substances (PFASs) and polycyclic aromatic hydrocarbon (PAH) metabolites, revealing a contamination gradient at the four stations. Furthermore, hepatosomatic index (HSI) and Fulton's condition factor (CF) were significantly lower in cod caged closest to the disposal site. Levels and activities of biomarker proteins, such as vitellogenin (Vtg), metallothionein (Mt), and biotransformation and oxidative stress enzymes, including cytochrome P450 1a and 3a (Cyp1a, Cyp3a), glutathione s-transferase (Gst) and catalase (Cat), were quantified in blood plasma and liver tissue. Hepatic Cat and Gst activities were significantly reduced in cod caged at the innermost stations in Kollevåg, indicating modulation of oxidative stress responses. However, these results contrasted with reduced hepatic lipid peroxidation. Significant increases in transcript levels were observed for genes involved in lipid metabolism (fasn and acly) in cod liver, while transcript levels of ovarian steroidogenic enzyme genes such as p450scc, cyp19, 3β-hsd and 20β-hsd showed significant station-dependent increases. Cyp1a and Vtg protein levels were however not significantly altered in cod caged in Kollevåg. Plasma levels of estradiol (E2) and testosterone (T) were determined by enzyme immunoassay (EIA) and showed elevated E2 levels, but only at the innermost station. We conclude that the bay of Kollevåg did not fullfill adequate environmental condition based on environmental quality standards (EQSs) for chemicals in coastal waters. Following a six weeks caging period, environmental contaminants accumulated in cod tissues and effects were observed on biomarker responses, especially those involved in reproductive processes in cod ovary., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Toxicity of weathered oil was investigated using Atlantic cod (Gadus morhua) larvae. A novel exposure system was applied to differentiate effects associated with dissolved and droplet oil with and without dispersant. After a 4-day exposure and subsequent 4-day recovery period, survival and growth were determined. Analytical data characterizing test oil composition included polyaromatic hydrocarbons (PAH) based on GC/MS and unresolved hydrocarbon classes obtained by two-dimensional chromatography coupled with flame ionization detection was used as input to an oil solubility model to calculate toxic units (TUs) of dissolved PAHs and whole oil, respectively. Critical target lipid body burdens derived from modeling characterizing the sensitivity of effect endpoints investigated were consistent across treatments and within the range previously reported for pelagic species. Individually measured PAHs captured only 3-11% of the TUs associated with the whole oil highlighting the limitations of traditional total PAH exposure metrics for expressing oil toxicity data., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Climate warming and harvesting affect the dynamics of species across the globe through a multitude of mechanisms, including distribution changes. In fish, migrations to and distribution on spawning grounds are likely influenced by both climate warming and harvesting. The Northeast Arctic (NEA) cod (Gadus morhua) performs seasonal migrations from its feeding grounds in the Barents Sea to spawning grounds along the Norwegian coast. The distribution of cod between the spawning grounds has historically changed at decadal scales, mainly due to variable use of the northern and southern margins of the spawning area. Based on historical landing records, two major hypotheses have been put forward to explain these changes: climate and harvesting. Climate could affect the distribution through, for example, spatial habitat shifts. Harvesting could affect the distribution through impacting the demographic structure. If demographic structure is important, theory predicts increasing spawner size with migration distance. Here, we evaluate these hypotheses with modern data from a period (2000-2016) of increasing temperature and recovering stock structure. We first analyze economic data from the Norwegian fisheries to investigate geographical differences in size of spawning fish among spawning grounds, as well as interannual differences in mean latitude of spawning in relation to changes in temperature and demographic parameters. Second, we analyze genetically determined fish sampled at the spawning grounds to unambiguously separate between migratory NEA cod and potentially smaller sized coastal cod of local origin. Our results indicate smaller spawners farther away from the feeding grounds, hence not supporting the hypothesis that harvesting is a main driver for the contemporary spawning ground distribution. We find a positive correlation between annual mean spawning latitude and temperature. In conclusion, based on contemporary data, there is more support for climate compared to harvesting in shaping spawning ground distribution in this major fish stock in the North Atlantic Ocean., (© 2018 John Wiley & Sons Ltd.)

Rapid climate change in the Northeast Atlantic and Arctic poses a threat to some of the world's largest fish populations. Impacts of warming and acidification may become accessible through mechanism-based risk assessments and projections of future habitat suitability. We show that ocean acidification causes a narrowing of embryonic thermal ranges, which identifies the suitability of spawning habitats as a critical life-history bottleneck for two abundant cod species. Embryonic tolerance ranges linked to climate simulations reveal that ever-increasing CO 2 emissions [Representative Concentration Pathway (RCP) 8.5] will deteriorate suitability of present spawning habitat for both Atlantic cod ( Gadus morhua ) and Polar cod ( Boreogadus saida ) by 2100. Moderate warming (RCP4.5) may avert dangerous climate impacts on Atlantic cod but still leaves few spawning areas for the more vulnerable Polar cod, which also loses the benefits of an ice-covered ocean. Emissions following RCP2.6, however, support largely unchanged habitat suitability for both species, suggesting that risks are minimized if warming is held "below 2°C, if not 1.5°C," as pledged by the Paris Agreement.

Food searching behaviour in a group of individually tagged 1-5 kg Atlantic cod Gadus morhua was studied in a set of three experiments in a sea cage with two underwater platforms, where restricted amounts of food was delivered several times per day during an acoustic training period. It took c. 1 week to train 20 naïve cod to associate low frequency (250 Hz) sound with food, whereas the training time was reduced to less than 2 days when 19 naïve G. morhua were accompanied with one trained fish. The fish formed a school that cruised between the platforms in search of food. Usually, there was one leader in the school, a fish that swam faster, arrived first at the platforms and visited the platforms more frequently than other members of the school. The leader spent more energy on swimming but also received more food and grew faster than the rest of the fish. At the start of the experiments, the leaders were not larger than the average fish but always among the leanest ones in the group. The study reveals how social learning can facilitate the acoustic training in adult G. morhua, information that may be useful in finding ways to aggregate valuable fish species for environmentally friendly fishing and ranching., (© 2018 The Fisheries Society of the British Isles.)

Crude oil accidentally spilled into the marine environment undergoes natural weathering processes that result in oil components being dissolved into the water column or present in particulate form as dispersed oil droplets. Oil components dissolved in seawater are typically considered as more bioavailable to pelagic marine organisms and the main driver of crude oil toxicity, however, recent studies indicate that oil droplets may also contribute. The adhesion of crude oil droplets onto the eggs of pelagic fish species may cause enhanced transfer of oil components via the egg surface causing toxicity during the sensitive embryonic developmental stage. In the current study, we utilized an oil droplet dispersion generator to generate defined oil droplets sizes/concentrations and exposed Atlantic cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) to investigate if the potential for dispersed oil droplets to adhere onto the surface of eggs was species-dependent. The influence of a commercial chemical dispersant on the adhesion process was also studied. A key finding was that the adhesion of oil droplets was significantly higher for haddock than cod, highlighting key differences and exposure risks between the two species. Scanning electron microscopy indicates that the differences in oil droplet adhesion may be driven by the surface morphology of the eggs. Another important finding was that the adhesion capacity of oil droplets to fish eggs is significantly reduced (cod 37.3%, haddock 41.7%) in the presence of the chemical dispersant., (Copyright © 2018 Elsevier B.V. All rights reserved.)

Predicting fish stock variations on interannual to decadal time scales is one of the major issues in fisheries science and management. Although the field of marine ecological predictions is still in its infancy, it is understood that a major source of multi-year predictability resides in the ocean. Here we show the first highly skilful long-term predictions of the commercially valuable Barents Sea cod stock. The 7-year predictions are based on the propagation of ocean temperature anomalies from the subpolar North Atlantic toward the Barents Sea, and the strong co-variability between these temperature anomalies and the cod stock. Retrospective predictions for the period 1957-2017 capture well multi-year to decadal variations in cod stock biomass, with cross-validated explained variance of over 60%. For lead times longer than one year the statistical long-term predictions show more skill than operational short-term predictions used in fisheries management and lagged persistence forecasts. Our results thus demonstrate the potential for ecosystem-based fisheries management, which could enable strategic planning on longer time scales. Future predictions show a gradual decline in the cod stock towards 2024., Competing Interests: The authors have declared that no competing interests exist.

The trophic link between cod (Gadus sp.) and capelin (Mallotus sp.) is important in many panarctic ecosystems. Since the early 2000s, the Northeast Arctic cod stock (G. morhua) in the Barents Sea has increased greatly, and the sea has been exceptionally warm. Such changes have potentially large effects on species distributions and overlap, which in turn could affect the strength of species interactions. Due to its high latitude location, the Barents Sea has strong seasonal variation in physical conditions and interactions. To study drivers of variation in cod-capelin overlap, we use data from two annual surveys run in winter and in autumn of 2004-2015. We first model winter and autumn spatial distributions of mature and immature cod and capelin. We then calculate overlap from model predictions on a grid with similar spatial resolution as the survey data. Our approach allowed us to interpret changes in overlap as species-specific effects of stock size and temperature, while accounting for sampling variation due to sampling time and depth. We found that during winter both species expanded their distribution in response to increased stock sizes, but how strongly and where the expansion occurred varied. The effect of temperature on distributions varied in space, and differed for cod and capelin and for different components of the two species. The results for autumn were clearer and more consistent. Both species expanded their distribution areas as their stock sizes increased. A positive effect of temperature was found in the north-eastern Barents Sea, where temperatures were lowest at the start of the study. Overlap increased and shifted north-eastwards during the study period and remained high despite a decline in the capelin stock. The increased overlap during autumn could mainly be attributed to the shift in cod distribution with increased cod stock biomass., Competing Interests: The authors have declared that no competing interests exist.

Trade-offs in energy allocation between growth, reproduction, and survival are at the core of life-history theory. While age-specific mortality is considered to be the main determinant of the optimal allocation, some life-history strategies, such as delayed or skipped reproduction, may be better understood when also accounting for reproduction costs. Here, we present a two-pool indeterminate grower model that includes survival and energetic costs of reproduction. The energetic cost sets a minimum reserve required for reproduction, while the survival cost reflects increased mortality from low postreproductive body condition. Three life-history parameters determining age-dependent energy allocation to soma, reserve, and reproduction are optimized, and we show that the optimal strategies can reproduce realistic emergent growth trajectories, maturation ages, and reproductive outputs for fish. The model predicts maturation phase shifts along the gradient of condition-related mortality and shows that increased harvesting will select for earlier maturation and higher energy allocation to reproduction. However, since the energetic reproduction cost sets limits on how early an individual can mature, an increase in fitness at high harvesting can only be achieved by diverting most reserves into reproduction. The model presented here can improve predictions of life-history responses to environmental change and human impacts because key life-history traits such as maturation age and size, maximum body size, and size-specific fecundity emerge dynamically.

In vitro cell culture methods are crucial for the isolation, purification and mass propagation of intracellular pathogens of aquatic organisms. Cell culture infection models can yield insights into infection mechanisms, aid in developing methods for disease mitigation and prevention, and inform commercial-scale cultivation approaches. This study details the establishment of a larval cell line (GML-5) from the Atlantic cod (Gadus morhua) and its use in the study of microsporidia. GML-5 has survived over 100 passages in 8 years of culture. The line remains active and viable between 8 and 21°C in Leibovitz-15 (L-15) media with 10% foetal bovine serum and exhibits a myofibroblast phenotype as indicated by immuno-positive results for vimentin, α-smooth muscle actin, collagen I and S-100 proteins, while being desmin-negative. GML-5 supports the infection and development of two microsporidian parasites, an opportunistic generalist (Anncaliia algerae) and cod-specific Loma morhua. Using GML-5, spore germination and proliferation of L. morhua was found to require exposure to basic pH and cool incubation temperatures (8°C), in contrast to A. algerae, which required no cultural modifications. Loma morhua-associated xenoma-like structures were observed 2 weeks postexposure. This in vitro infection model may serve as a valuable tool for cod parasitology and aquaculture research., (© 2018 John Wiley & Sons Ltd.)

Oil contamination is an environmental issue of great concern and the necessity for background studies and monitoring programs to continuously evaluate the levels of oil pollution is required. In this study, Atlantic cod (Gadus morhua) were exposed to dispersed crude oil for 1 and 4 weeks to simulate environmental contamination. Fractionated plasma samples were then analysed by tandem mass spectrometry. In total, 717 proteins were identified and 10 new protein biomarker candidates were found. The significant proteome changes were related to the immune response by alterations in the levels of specific immunoglobulins, alpha-2-macroglobulin and galectin-3-binding proteins. After 4 weeks of oil exposure, a lowered level of a NLRC3-like protein was also observed. The results from this study provide insight into the Atlantic cod plasma proteome and into the toxicological effects and potential response mechanisms of short and long-term exposure to crude oil., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Sustainability of commercial fisheries is best achieved when fishing gears are selective and have low impacts on bottom habitat. Pots (baited traps) are a fishing technology that typically has lower impacts than many other industrial gears. In this study we compared the efficiency of five models of pots (baited traps) designed to catch Atlantic cod (Gadus morhua) for use in Newfoundland and Labrador (NL)'s expanding cod fishery. We compared catch per unit effort (CPUE) and total lengths of cod across each pot type, as well as bycatch rates of each model. All pot types were successful at catching cod, but two models (the modified Newfoundland pot, and a four-entrance pot of our design) had highest CPUE. Specifically, we found that modifying Newfoundland pots increased their CPUE by 145% without a corresponding increase in bycatch. None of the pot types produced substantial amounts of bycatch. This study demonstrated that potting gear is an effective way to catch cod in NL, and that there is flexibility in which pot fishers can use, depending on the layout of their fishing vessel., Competing Interests: The authors have declared that no competing interests exist.

Ocean acidification (OA), the dissolution of excess anthropogenic carbon dioxide in ocean waters, is a potential stressor to many marine fish species. Whether species have the potential to acclimate and adapt to changes in the seawater carbonate chemistry is still largely unanswered. Simulation experiments across several generations are challenging for large commercially exploited species because of their long generation times. For Atlantic cod (Gadus morhua), we present first data on the effects of parental acclimation to elevated aquatic CO 2 on larval survival, a fundamental parameter determining population recruitment. The parental generation in this study was exposed to either ambient or elevated aquatic CO 2 levels simulating end-of-century OA levels (~1100 µatm CO 2 ) for six weeks prior to spawning. Upon fully reciprocal exposure of the F1 generation, we quantified larval survival, combined with two larval feeding regimes in order to investigate the potential effect of energy limitation. We found a significant reduction in larval survival at elevated CO 2 that was partly compensated by parental acclimation to the same CO 2 exposure. Such compensation was only observed in the treatment with high food availability. This complex 3-way interaction indicates that surplus metabolic resources need to be available to allow a transgenerational alleviation response to ocean acidification.

The semi-enclosed nature and estuarine characteristics, together with its strongly alternating bathymetry, make the Baltic Sea prone to much stronger interannual variations in the abiotic environment, than other spawning habitats of Atlantic cod (Gadus morhua). Processes determining salinity and oxygen conditions in the basins are influenced both by long term gradual climate change, e.g. global warming, but also by short-term meteorological variations and events. Specifically one main factor influencing cod spawning conditions, the advection of highly saline and well-oxygenated water masses from the North Sea, is observed in irregular frequencies and causes strong interannual variations in stock productivity. This study investigates the possibility to use the available hydrographic process knowledge to predict the annual spawning conditions for Eastern Baltic cod in its most important spawning ground, the Bornholm Basin, only by salinity measurements from a specific location in the western Baltic. Such a prediction could serve as an environmental early warning indicator to inform stock assessment and management. Here we used a hydrodynamic model to hindcast hydrographic property fields for the last 40+ years. High and significant correlations were found for months early in the year between the 33m salinity level in the Arkona Basin and the oxygen-dependent cod spawning environment in the Bornholm Basin. Direct prediction of the Eastern Baltic cod egg survival in the Bornholm Basin based on salinity values in the Arkona Basin at the 33 m depth level is shown to be possible for eggs spawned by mid-age and young females, which currently predominate the stock structure. We recommend to routinely perform short-term predictions of the Eastern Baltic cod spawning environment, in order to generate environmental information highly relevant for stock dynamics. Our statistical approach offers the opportunity to make best use of permanently existing infrastructure in the western Baltic to timely provide scientific knowledge on the spawning conditions of Eastern Baltic cod. Furthermore it could be a tool to assist ecosystem-based fisheries management with a cost-effective implementation by including the short term predictions as a simple indicator in the annual assessments.

The disappearance of larger individuals and the decrease in individual body condition suffered by Atlantic cod Gadus morhua in the eastern Baltic during the past two decades can be expected to affect the stock reproductive output. To investigate this, female G. morhua were collected during the spawning and pre-spawning period in 2015-2016. The current individual potential fecundity (F P ) of eastern Baltic G. morhua was estimated and analysed in relation to total length (L T ) and indices of nutritional status such as body condition (K) and hepato-somatic index (I H ) using generalized linear models. In addition, the current prevalence of atresia and its potential relation to K were investigated. Moreover, a calibration curve to estimate F P from oocyte diameter, based on the autodiametric oocyte counting method, was established for the first time for eastern Baltic G. morhua and can be used for future fecundity studies on this stock. The results showed that F P was mainly positively related to fish length, but K and I H also contributed significantly to the variation in F P . The model predicted that fish with K = 1·2 have a F P 51% higher than fish of the same L T with K = 0·8. The prevalence of fecundity regulation by atresia was 5·8%, but it was found only in fish in the pre-spawning maturity stage and with low K. Temporal changes in biological features such as the length composition and individual body condition of eastern Baltic G. morhua, should be accounted for when estimating stock reproductive potential., (© 2018 The Fisheries Society of the British Isles.)

Most fishes and crustaceans respond to light, and artificial light sources may therefore be an efficient stimulus to manipulate behaviours in aquatic animals. It has been hypothesised that the catch efficiency of pots could be increased if prey, for example krill, can be attracted into the pots providing a visual stimulus and a source of live bait. To find which light characteristics are most attractive to krill, we tested the effects of light intensity and wavelength composition on Northern krill's (Meganyctiphanes norvegica) behavioural response to an artificial light source. The most attractive individual wavelength was 530 nm (green light), while broadband (425-750 nm) white light was an equally attractive light source. The intensity of the emitted light did not appear to have a direct effect on attraction to the light source, however it did significantly increase swimming activity among the observed krill. The most promising light stimuli for krill were tested to determine whether they would have a repulsive or attractive effect on cod (Gadus morhua); These light stimuli appeared to have a slightly repulsive, but non-significant, effect on cod. However, we suggest that a swarm of krill attracted to an artificial light source may produce a more effective visual stimulus to foraging cod.

The visual system is for many fishes essential in guiding behaviors, such as foraging, predator avoidance and mate choice. The marine environment is characterized by large spatio-temporal fluctuations in light intensity and spectral composition. However, visual capabilities are restricted by both space limitations set by eye size and by the genomic content of light-absorbing opsin genes. The rich array of visual opsins in teleosts may be used differentially to tune vision towards specific needs during ontogeny and to changing light. Yet, to what extent visual plasticity is a pre-programmed developmental event, or is triggered by photic environment, is unclear. Our previous studies on Atlantic cod revealed an evolutionary genomic loss of UV-sensitive sws1 and red-sensitive lws opsin families, while blue-sensitive sws2 and green-sensitive rh2 opsins had duplicated. The current study has taken an opsin expression approach to characterize visual plasticity in cod towards different spectral light during the larval stage, to maturation and extreme seasonal changes in the Barents Sea. Our data suggest that opsin plasticity in cod larvae is controlled by developmental programme rather than immediate light environment. The lack of expressional changes during maturation suggests a less important role for visual modulation related to mate choice. Although no seasonal effects on visual opsins were detected in migratory Northeast Arctic cod, the expressed opsin subset differed from the more stationary Norwegian coastal cod described in previous studies. Interestingly, these data provide the first indications of a population difference in actively used visual opsins associated with cod ecotypes., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

In the current study activity and latency to explore, as well as the correlation of these traits, were examined in individually marked juvenile Gadus morhua at 7, 10 and 13° C. It was concluded that individual rank order of both traits was maintained across temperature but that the level of change differed between individuals., (© 2017 The Fisheries Society of the British Isles.)

Productivity of marine fish stocks is known to be affected by environmental and ecological drivers, and global climate change is anticipated to alter recruitment success of many stocks. While the direct effects of environmental drivers on fish early life stage survival can be quantified experimentally, indirect effects in marine ecosystems and the role of adaptation are still highly uncertain. We developed an integrative model for the effects of ocean warming and acidification on the early life stages of Atlantic cod in the Barents Sea, termed SCREI (Simulator of Cod Recruitment under Environmental Influences). Experimental results on temperature and CO 2 effects on egg fertilization, egg and larval survival and development times are incorporated. Calibration using empirical time series of egg production, temperature, food and predator abundance reproduces age-0 recruitment over three decades. We project trajectories of recruitment success under different scenarios and quantify confidence limits based on variation in experiments. A publicly accessible web version of the SCREI model can be run under www.oceanchange.uni-bremen.de/;SCREI. Severe reductions in average age-0 recruitment success of Barents Sea cod are projected under uncompensated warming and acidification toward the middle to end of this century. Although high population stochasticity was found, considerable rates of evolutionary adaptation to acidification and shifts in organismal thermal windows would be needed to buffer impacts on recruitment. While increases in food availability may mitigate short-term impacts, an increase in egg production achieved by stock management could provide more long-term safety for cod recruitment success. The SCREI model provides a novel integration of multiple driver effects in different life stages and enables an estimation of uncertainty associated with interindividual and ecological variation. The model thus helps to advance toward an improved empirical foundation for quantifying climate change impacts on marine fish recruitment, relevant for ecosystem-based assessments of marine systems under climate change., (© 2017 John Wiley & Sons Ltd.)

Intraspecific diversity is central to the management and conservation of exploited species, yet knowledge of how this diversity is distributed and maintained in the genome of many marine species is lacking. Recent advances in genomic analyses allow for genome-wide surveys of intraspecific diversity and offer new opportunities for exploring genomic patterns of divergence. Here, we analysed genome-wide polymorphisms to measure genetic differentiation between an offshore migratory and a nonmigratory population and to define conservation units of Atlantic Cod (Gadus morhua) in coastal Labrador. A total of 141 individuals, collected from offshore sites and from a coastal site within Gilbert Bay, Labrador, were genotyped using an ~11k single nucleotide polymorphism array. Analyses of population structure revealed strong genetic differentiation between migratory offshore cod and nonmigratory Gilbert Bay cod. Genetic differentiation was elevated for loci within a chromosomal rearrangement found on linkage group 1 (LG1) that coincides with a previously found double inversion associated with migratory and nonmigratory ecotype divergence of cod in the northeast Atlantic. This inverted region includes several genes potentially associated with adaptation to differences in salinity and temperature, as well as influencing migratory behaviour. Our work provides evidence that a chromosomal rearrangement on LG1 is associated with parallel patterns of divergence between migratory and nonmigratory ecotypes on both sides of the Atlantic Ocean., (© 2017 John Wiley & Sons Ltd.)