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Native to the Ponto-Caspian region, the benthic round goby (Neogobius melanostomus) has invaded several European inland waterbodies as well as the North American Great Lakes and the Baltic Sea. The species is capable of reaching very high densities in the invaded ecosystems, with not only evidence for significant food-web effects on the native biota and habitats, but also negative implications to coastal fishers. Although generally considered a coastal species, it has been shown that round goby migrate to deeper areas of the Great Lakes and other inland lakes during the cold season. Such seasonal movements may create new spatio-temporal ecosystem consequences in invaded systems. To seek evidence for seasonal depth distribution in coastal marine habitats, we compiled all available catch data for round goby in the Baltic Sea since its invasion and until 2017. We furthermore related the depths at capture for each season with the ambient thermal environment. The round goby spend autumn and winter at significantly deeper and offshore areas compared to spring and summer months; few fish were captured at depths 25 m. The thermal conditions at which round goby were caught varied significantly between seasons, being on average 18.3 °C during summer, and dropping to a low 3.8 °C during winter months. Overall, the fish sought the depths within each season with the highest possible temperatures. The spatial distribution of the round goby substantially overlaps with that of its main and preferred prey (mussels) and with that of its competitor for food (flatfish), but only moderately with the coastal predatory fish (perch), indicating thereby very complex trophic interactions associated with this invasion. Further investigations should aim at quantifying the food web consequences and coupling effects between different habitats related to seasonal migrations of the round goby, both in terms of the species as a competitor, predator and prey.

The round goby (Neogobius melanostomus) is a fish native to the Ponto-Caspian region that is highly invasive through freshwater and brackish habitats in northern Europe and North America. Individual behavioural variation appears to be an important factor in their spread, for example a round goby’s personality traits can influence their dispersal tendency, which may also produce variation in the behavioural composition of populations at different points along their invasion fronts. To further analyze the drivers of behavioural variation invasive round goby populations, we focused on two populations along the Baltic Sea invasion front with closely comparable physical and community characteristics. Specifically, this study measured personality within a novel environment and predator response context (i.e., boldness), and directly analyzed links between individuals’ personality traits and their physiological characteristics and stress responses (i.e., blood cortisol and lactate, brain neurotransmitters). In contrast to previous findings, the more recently established population had similar activity levels but were less bold in response to a predator cue than the older population, which suggests that behavioural compositions within our study populations may be more driven by local environmental conditions rather than being a result of personality-biased dispersal. Furthermore, we found that both populations showed similar physiological stress responses, and there also appeared to be no detectable relationship between physiological parameters and behavioural responses to predator cues. Instead, body size and body condition were important factors influencing individual behavioural responses. Overall, our results reinforce the importance of boldness traits as a source of phenotypic variation in round goby populations in the Baltic Sea, but that the physiological mechanisms underpinning personality trait variation in these populations remain unclear.

Species invasions often occur on coasts and estuaries where abiotic conditions vary, e.g. salinity, temperature, runoff etc. Successful establishment and dispersal of non-indigenous species in many such systems are poorly understood, partially since the species tend to show genetic and ecological plasticity at population level towards many abiotic conditions, including salinity tolerance. Plasticity may be driven by shifting expression of heat shock proteins such as Hsp70, which is widely recognized as indicator of physical stress. In this study, we developed a qPCR assay for expression of the hsp70 gene in the invasive round goby (Neogobius melanostomus) and tested the expression response of fish collected from a brackish environment in the western Baltic Sea to three different salinities, 0, 10 and 30. hsp70 expression was highest in fresh water, indicating higher stress, and lower at brackish (ambient condition for the sampled population) and oceanic salinities, suggestive of low stress response to salinities above the population’s current distribution. The highest stress in fresh water was surprising since populations in fresh water exist, e.g. large European rivers and Laurentian Great Lakes. The results have implications to predictions for the species’ plasticity potential and possible range expansion of the species into other salinity regimes.

During the 2010s, Atlantic cod Gadus morhua L. in the eastern Baltic Sea experienced increasing infection loads of the parasitic nematode Contracaecum osculatum (Rudolphi) in their livers. Starting in 2021, a mandatory part of the routine sampling protocol on Baltic monitoring surveys is to assign a liver category to individual cod livers, based on the number of nematodes visible on the liver surface, to follow spatiotemporal changes in nematode infection loads. The validity of the liver category method has never been evaluated. Based on data from 642 cod livers, the method was verified and found to be a good predictor of the total number of nematodes. Moreover, the probability of cod being in a critical condition increased with the parasite load. In addition to their direct applicability to Baltic cod, the present findings may inspire others working with disease in fish stocks to include parasite monitoring.

Invasive species exert negative impacts on biodiversity and ecosystems on a global scale, which may be enhanced in the future by climate change. Knowledge of how invasive species respond physiologically and behaviorally to novel and changing environments can improve our understanding of which traits enable the ecological success of these species, and potentially facilitate mitigation efforts. We examined the effects of acclimation to temperatures ranging from 5 to 28°C on aerobic metabolic rates, upper temperature tolerance (critical thermal maximum, CTmax), as well as temperature preference (Tpref) and avoidance (Tavoid) of the round goby (Neogobius melanostomus), one of the most impactful invasive species in the world. We show that round goby maintained a high aerobic scope from 15 to 28°C; that is, the capacity to increase its aerobic metabolic rate above that of its maintenance metabolism remained high across a broad thermal range. Although CTmax increased relatively little with acclimation temperature compared with other species, Tpref and Tavoid were not affected by acclimation temperature at all, meaning that round goby maintained a large thermal safety margin (CTmax−Tavoid) across acclimation temperatures, indicating a high level of thermal resilience in this species. The unperturbed physiological performance and high thermal resilience were probably facilitated by high levels of phenotypic buffering, which can make species readily adaptable and ecologically competitive in novel and changing environments. We suggest that these physiological and behavioral traits could be common for invasive species, which would only increase their success under continued climate change., Summary: Phenotypic buffering of physiology and behavior, coupled with large thermal safety margins, may be competitively advantageous for the invasive fish round goby, and enhance its invasive potential with climate change.

For externally fertilising organisms in the aquatic environment, the abiotic fertilisation medium can be a strong selecting force. Among bony fishes, sperm are adapted to function in a narrow salinity range. A notable exception is the family Gobiidae, where several species reproduce across a wide salinity range. The family also contains several wide-spread invasive species. To better understand how these fishes tolerate such varying conditions, we measured sperm performance in relation to salinity from a freshwater and a brackish population within their ancestral Ponto-Caspian region of the round goby, Neogobius melanostomus. These two ancestral populations were then compared to nine additional invaded sites across northern Europe, both in terms of their sperm traits and by using genomic SNP markers. Our results show clear patterns of ancestral adaptations to freshwater and brackish salinities in their sperm performance. Population genomic analyses show that the ancestral ecotypes have generally established themselves in environments that fit their sperm adaptations. Sites close to ports with intense shipping show that both outbreeding and admixture can affect the sperm performance of a population in a given salinity. Rapid adaptation to local conditions is also supported at some sites. Historical and contemporary evolution in the traits of the round goby sperm cells is tightly linked to the population and seascape genomics as well as biogeographic processes in these invasive fishes. Since the risk of a population establishing in an area is related to the genotype by environment match, port connectivity and the ancestry of the round goby population can likely be useful for predicting the species spread.For externally fertilising organisms in the aquatic environment, the abiotic fertilisation medium can be a strong selecting force. Among bony fishes, sperm are adapted to function in a narrow salinity range. A notable exception is the family Gobiidae, where several species reproduce across a wide salinity range. The family also contains several wide-spread invasive species. To better understand how these fishes tolerate such varying conditions, we measured sperm performance in relation to salinity from a freshwater and a brackish population within their ancestral Ponto-Caspian region of the round goby, Neogobius melanostomus. These two ancestral populations were then compared to nine additional invaded sites across northern Europe, both in terms of their sperm traits and by using genomic SNP markers. Our results show clear patterns of ancestral adaptations to freshwater and brackish salinities in their sperm performance. Population genomic analyses show that the ancestral ecotypes have generally established themselves in environments that fit their sperm adaptations. Sites close to ports with intense shipping show that both outbreeding and admixture can affect the sperm performance of a population in a given salinity. Rapid adaptation to local conditions is also supported at some sites. Historical and contemporary evolution in the traits of the round goby sperm cells is tightly linked to the population and seascape genomics as well as biogeographic processes in these invasive fishes. Since the risk of a population establishing in an area is related to the genotype by environment match, port connectivity and the ancestry of the round goby population can likely be useful for predicting the species spread.

Aim: The few fish species able to reproduce across wide osmotic ranges either plastically acclimate sperm performance to, or are locally adapted to, different salinities. The invasive round goby (Neogobius melanostomus) is spreading in Eurasia and the Americas, into both fresh and brackish water. We aim to understand if reproduction in different salinities is affected by an ability to acclimate. Location: Brackish and freshwater systems of northern Europe and the Baltic Sea. Methods: We cross-exposed round gobies of freshwater and brackish origin to 0 and 16 practical salinity units (PSU), and the fish were given nest boxes in which to spawn. After 4 weeks, we measured their sperm performance in both 0 and 16 PSU; fertilization success of each egg clutch was measured through visual analysis of eggs. Clutches were split and allowed to develop in both 0 and 16 PSU salinity, and reproductive success (zygote development) was measured 20 ± 1 days later. Responses were analysed using generalized mixed models. Results: After a month, the fish showed no plasticity in sperm performance to their acclimation salinity, regardless of their origin. Sperm velocity was highest in the salinity similar to the males’ origin. Significantly lower fertilization success was measured for individuals that reproduced outside their salinity of origin despite recurring spawning events in all treatment groups. Among fertilized eggs, zygote development was similar regardless of salinity treatment of either eggs or parents. Main Conclusions: Short-term acclimation to new salinities does not affect sperm performance in the round goby. Alternative hypotheses such as local adaption should be further investigated. Limits to the species’ reproductive success, and therefore invasion processes, are likely dependent on environment-phenotype matches. Fish of brackish origin spawned successfully in freshwater, pointing to an increased risk of introducing populations of brackish origin into freshwater.

This study examines the impact of boldness on foraging competition of the highly invasive round goby Neogobius melanostomus Pallas 1815. Individual risk tolerance, or boldness, was measured as the time to resume movement after a simulated predation strike. Fish that resumed movement faster were categorized as “bold”, fish that took more time to resume movement were categorized as “shy”, and those who fell in between these two categories were determined to have “intermediate” boldness. Competitive impacts of boldness in N. melanostomus were determined in a laboratory foraging experiment in which interspecific (juvenile Atlantic cod Gadus morhua Linnaeus 1758) and intraspecific (intermediate N. melanostomus) individuals were exposed to either bold or shy N. melanostomus competitors. G. morhua consumed fewer prey when competing with bold N. melanostomus than when competing with shy N. melanostomus, whereas intermediately bold N. melanostomus foraging was not affected by competitor boldness. Bold and shy N. melanostomus consumed similar amounts of prey, and the number of interactions between paired fish did not vary depending on the personality of N. melanostomus individuals. Hence, intraspecific foraging competition was not found to be personality dependent. This study provides evidence that individual differences in boldness can mediate competitive interactions in N. melanostomus, however results also show that competition is also governed by other mechanisms which require further study.

The nutritional condition; the energy turnover; the body, liver and plasma composition; and the digestive organ masses were evaluated in relation to varying infection densities with liver worm in Eastern Baltic cod. We show that fish with high infection load with this parasitic nematode have severely compromised physiological condition and poor health status., Establishing relationships between parasite infection and physiological condition of the host can be difficult and therefore are often neglected when describing factors causing population declines. Using the parasite–host system between the parasitic nematode Contracaecum osculatum and the Eastern Baltic cod Gadus morhua, we here shed new light on how parasite load may relate to the physiological condition of a transport host. The Eastern Baltic cod is in distress, with declining nutritional conditions, disappearance of the larger fish, high natural mortality and no signs of recovery of the population. During the latest decade, high infection levels with C. osculatum have been observed in fish in the central and southern parts of the Baltic Sea. We investigated the aerobic performance, nutritional condition, organ masses, and plasma and proximate body composition of wild naturally infected G. morhua in relation to infection density with C. osculatum. Fish with high infection densities of C. osculatum had (i) decreased nutritional condition, (ii) depressed energy turnover as evidenced by reduced standard metabolic rate, (iii) reduction in the digestive organ masses, and alongside (iv) changes in the plasma, body and liver composition, and fish energy source. The significantly reduced albumin to globulin ratio in highly infected G. morhua suggests that the fish suffer from a chronic liver disease. Furthermore, fish with high infection loads had the lowest Fulton’s condition factor. Yet, it remains unknown whether our results steam from a direct effect of C. osculatum, or because G. morhua in an already compromised nutritional state are more susceptible towards the parasite. Nevertheless, impairment of the physiological condition can lead to reduced swimming performance, compromising foraging success while augmenting the risk of predation, potentially leading to an increase in the natural mortality of the host. We hence argue that fish–parasite interactions must not be neglected when implementing and refining strategies to rebuild deteriorating populations.

Passive integrated transponder (PIT) tags are commonly used to assess fish movement for use in fisheries management. Here, we investigated physiological and behavioral effects of tagging on sandeels (Ammodytes tobianus) using PIT tags constituting 2.1 ± 0.9% of their body weight. Swimming stamina (RSS), calculated as time spent swimming against the current relative to total swimming time, and tail beat frequency were compared between tagged and untagged fish as was blood hematocrit levels at 7, 14, and 42 d post-tagging. Survival and tag retention were also documented at 14, 42, and 84 d (via x-rays and dissections). RSS was not different between tagged and untagged fish with means (± SD) of 60 ± 9% and 61 ± 12%. Tail beat frequency was not different between tagged and untagged fish at 2.8 ± 0.3 and 3.0 ± 0.4 beats s−1 for tagged and untagged fish, respectively. Likewise, hematocrit was not affected by tagging and levels were between 21–26% for both groups. Survival rates were high and did not differ between groups (96% for tagged and 99% untagged fish). Tag retention was 100%. X-rays and dissections did not reveal any signs of tag movement at 14–84 d, and there was no difference between relative positions of the tags. None of the tags were encapsulated in the body cavity after 14 d, whereas 40 and 56% of the tags were encapsulated in a thin tissue membrane between the intestine and kidney after 42 and 84 d, respectively. After 14 d all incisions had healed with only minor or no signs of the tag insertion site. Collectively, these data provide substantial evidence for the possibility of conducting large-scale tagging studies on this species in the field.

Larvae of the eye fluke, Diplostomum, emerge from snails and infect fish by penetrating skin or gills, then move to the lens where they may impair the vision of the fish. For the fluke to reproduce, a bird must eat the infected fish, and it has been suggested that they therefore actively manipulate the fish’s behaviour to increase the risk of predation. We found that round gobies Neogobius melanostomus, a species that was recently introduced to the Kalmar Sound of the Baltic Sea, had an eye fluke prevalence of 90–100%. We investigated how the infection related to behavioural variation in round gobies. Our results showed that the more intense the parasite-induced cataract, the weaker the host’s response was to simulated avian attack. The eye flukes did not impair other potentially important anti-predator behaviours, such as shelter use, boldness and the preference for shade. Our results are in accordance with the suggestion that parasites induce changes in host behaviour that will facilitate transfer to their final host.

Many aquatic ecosystems are invaded by alien species, often with severe implications for native species and associated fisheries. Round goby Neogobius melanostomus is of Ponto-Caspian origin and has established large populations in the Baltic Sea. Here, round goby often feed on blue mussel Mytilus edulis, which may affect the invaded ecosystems. Experimental data were modelled and showed that round goby up to c. 15 cm of total length (TL) feed on mussels

Reefs are structurally complex habitats that are degraded in numerous coastal areas. Structural complexity is often associated with elevated fish abundance, and recent studies have indicated that such structural complexity (e.g. reefs) not only acts as a fish aggregator, but also increases fish production. The objective of this study was to advance this knowledge by investigating if an underlying mechanism of the observed productivity is related to reduced metabolic rates (proxy for energy use) of fish in reef habitats. Using juvenile Atlantic cod Gadus morhua, we tested the hypothesis that fish energy use differs between fish occupying stone reef and sand bottom habitats. Metabolic rate (MO2) was estimated using intermittent flow respirometry in simulated stone reef and sand bottom habitats over 24 h. Results revealed that G. morhua in the stone reef habitat exhibited significantly reduced accumulated MO2 compared to G. morhua in the sand bottom habitat. Likewise, there was a tendency for lower mean standard metabolic rates of the fish in stone reefs, although this pattern was not statistically significant. There are many mechanisms that may underpin elevated productivity in structurally complex habitats such as reefs, including better access to shelter and increased food availability. Our study adds to these mechanisms by showing that G. morhua save energy when occupying stone reefs as compared to sandy bottoms, energy which may be allocated to somatic and gonadal growth.

Originating from the Ponto-Caspian region, the invasive round goby has since 1990 spread in the Baltic Sea region, now with established populations throughout the area. The fish is a strong competitor for food, shelter and nesting sites, and has a high tolerance towards a broad range of temperatures and salinities. In many places, it exerts negative effects on the local fish fauna and benthic community, and its presence may compromise descriptors of Good Environmental Status under the Marine Strategy Framework Directive. Furthermore, Marine protected areas, constructed to protect valuable and threatened species or habitats, may be compromised by its presence. Notably however, round goby is also an emerging prey for larger fish, marine mammals and birds, thus providing positive effects to the invaded ecosystems. In some areas, it also supports fisheries, adding positively to society.

The round goby (Neogobius melanostomus; Pallas, 1814) is one of the most widespread invasive fish species. It originates from the Black Sea and the Sea of Azov but has un-intentionally been introduced to fresh and brackish water bodies on both sides of the Atlantic. Small-scale fishermen catch large amounts of the invasive round goby as the species continues to spread and populate new areas of the Baltic Sea. We were concerned with the possibilities for a “use and reduce” solution to the round goby invasion. However, to capture the interest of restaurants, supermarkets, and distributers, more information about seasonality in product supply and nutritional quality is needed. We therefore studied a round goby bycatch fishery in the Western Baltic Sea (Denmark) and describe the seasonal dynamics in catch rates, size distribution, condition factor, lipid and protein content, fatty acid profile, and meat texture. We found that while the catch rates peaked in spring, lipid content and meat texture improved significantly, later in the year. Protein content also peaked in late summer and fall, but it was only marginally higher compared to spring. We compare the results to relevant species and conclude that round goby is a relatively lean fish all year (only slightly fatter than for example cod). The firmness of the meat in fall, resembles farmed trout and it has a highly favorable fatty acid composition, reflected by a low n-6 to n-3 ratio. The main bottleneck related to the introduction of round goby to the consumer market is likely going to be the small size.

Fish serve as transport hosts to a range of parasites, with potential negative effects on fish health. In the Baltic Sea, the grey seal Halichoerus grypus (Fabricius) population has increased markedly since the early 2000s. H. grypus is the main final host to the liver worm Contracaecum osculatum (Rudolphi, 1802), a parasitic nematode to which cod Gadus morhua (Linnaeus) is one of several transport hosts. Recent investigations have shown a marked increase in prevalence and abundance of infection of this parasite in livers of G. morhua inhabiting the central Baltic Sea. Yet no recent knowledge exists on levels of C. osculatum infection in G. morhua in adjacent areas. We investigated spatial differences in prevalence and abundance of this parasitic nematode in livers of G. morhua, covering a transect consisting of 9 areas from the Skagerrak to the eastern part of the central Baltic Sea. We further provide survey data of local abundances of H. grypus and harbor seal Phoca vitulina (Linnaeus) throughout this transect. Prevalence and abundance of C. osculatum sensu stricto in G. morhua livers differed significantly between east and west, with highest levels of infection occurring in the low-salinity central Baltic areas. Highly infected fish in the east had significantly lower condition factor than their westerly, less infected conspecifics. Spatial differences in local seal abundance and seal species, salinity and feeding ecology may explain the observed differences in C. osculatum infection between eastern and western G. morhua

Round goby Neogobius melanostomus is currently one of the most wide-ranging invasive fish species in Europe and North America. The present study demonstrates how the distribution of round goby has expanded from 2008 to 2013 at a rate of about 30 km yr−1 along the Danish coastline in the western Baltic Sea. Further analyses showed that fish from an establishedhigh-density round goby population were slow-growing and displayed poorer condition (weight at age and hepatosomatic index) compared to fish sampled from recently invaded locations (i.e. at the forefront of the distribution range). The established population revealed a broad age distribution and a 1:1 gender ratio, while fish from a recently invaded site were primarily of intermediateages with a male-biased gender ratio. Otolith analyses suggested that the oldest individuals from the recently invaded area experienced superior growth conditions only in the most recent years, suggesting immigration into the area as adults. Our results suggest that intraspecific competition for food may cause continued dispersal of the species and that population demographics likelyrelate to invasion history

The eastern Baltic (EB) cod (Gadus morhua) stock was depleted and overexploited for decades until the mid-2000s, when fishing mortality rapidly declined and biomass started to increase, as shown by stock assessments. These positive developments were partly assigned to effective management measures, and the EB cod was considered one of the most successful stock recoveries in recent times. In contrast to this optimistic view, the analytical stock assessment failed in 2014, leaving the present stock status unclear. Deteriorated quality of some basic input data for stock assessment in combination with changes in environmental and ecological conditions has led to an unusual situation for cod in the Baltic Sea, which poses new challenges for stock assessment and management advice. A number of adverse developments such as low nutritional condition and disappearance of larger individuals indicate that the stock is in distress. In this study, we (i) summarize the knowledge of recent changes in cod biology and ecosystem conditions, (ii) describe the subsequent challenges for stock assessment, and (iii) highlight the key questions where answers are urgently needed to understand the present stock status and provide scientifically solid support for cod management in the Baltic Sea.

Vertical migration is the most widespread migration in the aquatic world, yet the mechanisms limiting the extent of this behavior are largely unknown. In the Baltic Sea, some Atlantic cod Gadus morhua perform vertical foraging migrations into severely hypoxic demersal zones where aerobic metabolism is insufficient to cover energy requirements. After foraging, the fish return to better oxygenated waters for physiological recovery and digestion. To test the influence of phenotypic variation in aerobic scope (AS; the difference between the maximum and the minimum metabolic rate) on the capacity to migrate into severely hypoxic zones, we incorporated AS into a state-dependent individual-based model simulating vertical foraging migrations of G. morhua. We found little effect of individual variation in AS on the capacity for vertical migration when the zone used for physiological recovery was normoxic. In contrast, when there was moderate hypoxia (30% air saturation, O-2sat) in the zone used for physiological recovery, the high AS phenotype had a clear advantage because it could forage 3-4 times longer in the severely hypoxic (16% O-2sat, i.e. below the threshold for aerobic metabolism of the species) demersal zone compared to the low AS phenotype. Thus, phenotypic variation in AS is only important when there is moderate hypoxia in the zone used for physiological recovery, suggesting that the influence of AS variation on the capacity for vertical migration is context dependent. We propose that elevated AS may be evolutionarily favorable when hypoxia prevails in the water column.

Jakobshavn Glacier, west Greenland, has responded to temperature changes in Ilulissat Icefjord, into which it terminates. This study collected hydrographic observations inside Ilulissat Icefjord and from adjacent Disko Bay between 2001 and 2014. The warmest deep Disko Bay waters were blocked by the entrance sill and did not reach Jakobshavn Glacier. In the fjord basin, the summer mean temperature was 2.8°C from 2009 to 2013, excluding 2010, when it was 1°C cooler. Despite this variability, summer potential densities in the basin were in the narrow range of 27.20 ≤ σθ ≤ 27.31 kg m−3, and basin water properties matched those of Disko Bay in this layer each summer. This relation has likely held since at least 1980. Basin waters from 2009 and 2011–13 were therefore similar to those in 1998/99, when Jakobshavn Glacier began to retreat, while basin waters in 2010 were as cool as in the 1980s. The 2010 basin temperature anomaly was advected into Disko Bay, not produced by local atmospheric variability.This anomaly also shows that Ilulissat Icefjord basin waters were renewed annually or faster. Time series fragments inside the fjord did not capture the 2010 anomaly but show that the basin temperatures varied little subannually, outside of summer. Fjord velocity profiles from summer 2013 implied a basin renewal time scale of about 1 month. In model simulations of the fjord circulation, subglacial discharge from Jakobshavn Glacier could drive renewal of the fjord basin over a single summer, while baroclinic forcing from outside the fjord could not, because of the sill at the mouth.

Changes in ambient temperature affect the physiology and metabolism and thus the distribution of fish. In this study we used intermittent flow respirometry to determine the effect of temperature (2, 5, 10, 15 and 20°C) and wet body mass (BM) (~30-460g) on standard metabolic rate (SMR, mgO2h(-1)), maximum metabolic rate (MMR, mgO2h(-1)) and metabolic scope (MS, mgO2h(-1)) of juvenile Atlantic cod. SMR increased with BM irrespectively of temperature, resulting in an average scaling exponent of 0.87 (0.82-0.92). Q10 values were 1.8-2.1 at temperatures between 5 and 15°C but higher (2.6-4.3) between 2 and 5°C and lower (1.6-1.4) between 15 and 20°C in 200 and 450g cod. MMR increased with temperature in the smallest cod (50g) but in the larger cod MMR plateaued between 10, 15 and 20°C. This resulted in a negative correlation between the optimal temperature for MS (Topt) and BM, Topt being respectively 14.5, 11.8 and 10.9°C in a 50, 200 and 450g cod. Irrespective of BM cold water temperatures resulted in a reduction (30-35%) of MS whereas the reduction of MS at warm temperatures was only evident for larger fish (200 and 450g), caused by plateauing of MMR at 10°C and above. Warm temperatures thus seem favourable for smaller (50g) juvenile cod, but not for larger conspecifics (200 and 450g).

We report an unexpected observation of lesser sandeel Ammodytes marinus foraging on juveniles and late larval stages of the same species. This recording sheds new light on the cannibalistic and piscivorous capacity of forage fish and raises a number of questions about the role of forage fish in marine food webs. In 2012 and 2013 the stomachs of 748 sandeels from 36 different commercial sandeel hauls in the central North Sea were opened. 9% of these stomachs contained late stage sandeel larvae. In order to better understand the cannibalistic nature of sandeels, we made a detailed analysis of another 450 sandeels from a single haul with a high frequency of apparent cannibals. One-third of the stomachs contained a minimum of one young sandeel (mean length 2.7 cm; max. length 4.9 cm), 10 percent contained 5 or more, and one stomach contained 18. Analyses of sample DNA confirmed that predator and prey were conspecifics. Larger specimens were more likely to be cannibals. However, among cannibals the specific sandeel larvae consumption was independent of cannibal size. We argue that this piscivorous cannibalistic behaviour may not only be a key factor in explaining recruitment fluctuations in North Sea sandeel stocks, but it may also add a new element to the complexity of energy flow in marine food chains

The deep-water flatfish Greenland halibut Reinhardtius hippoglossoides (Walbaum) is common along the West Greenland coast. In the northwestern fjords, Greenland halibut is an important socio-economic resource for the Greenland community, but due to the deep and partly ice-covered environment, very little is known about its behavior and habitat characteristics. We tagged adult Greenland halibut in the waters off Ilulissat with electronic data storage tags that collected information on depth, temperature, and time. Although clear differences between individuals in migration and vertical behavior were present, we discovered a consistent seasonal migration from the relatively shallow-water Disko Bay area into the deep waters of the Ilulissat Icefjord, where the fish resided in the winter months before returning to Disko Bay. Vertical activity was pronounced at both locations, with fish covering vertical distances of up to 100 m within 15 min. During the winter months, the fish experienced temperatures between ca. 0 and 4°C, with most experiencing temperatures of 2 to 3°C. Irrespective of year and quarter of the year, the fish experienced warmer water and a broader range of temperatures when resident in Disko Bay (mean range 2.6°C) than when resident in the ice fjord (mean range 1.4°C). Using the tagged halibut as a 'live tool,' we show that parts of the ice fjord are hundreds of meters deeper than pre- viously thought. We also document the first seawater temperature measurements made beneath the Jakobshavn Isbrae outlet glacier, revealing a positive relationship between depth and temper- ature for the upper 600 m and a between-year variation in temperatures beneath the ice sheet in 2001, 2002, and 2003.

The parasitic copepod Lernaeocera branchialis had a negative effect on cardiorespiratory function in Atlantic cod Gadus morhua such that it caused pronounced cardiac dysfunction with irregular rhythm and reduced stroke amplitude compared with uninfected fish. In addition, parasite infection depressed the postprandial cardiac output and oxygen consumption.

Atlantic cod Gadus morhua experienced oxygen deficit ( D O 2 ) when exposed to oxygen levels below their critical level (c. 73% of pcrit ) and subsequent excess post-hypoxic oxygen consumption (CEPHO ) upon return to normoxic conditions, indicative of an oxygen debt. The mean ± s.e. CEPHO : D O 2 was 6·9 ± 1·5, suggesting that resorting to anaerobic energy production in severe hypoxia is energetically expensive.

Non-indigenous species (NIS) can impact marine biodiversity and ecosystem structure and function. Once introduced into a new region, secondary dispersal is limited by the physiology of the organism in relation to the ambient environment and by complex interactions between a suite of ecological factors such as presence of predators, competitors, and parasites. Early prediction of dispersal potential and future ‘area of impact’ is challenging, but also a great asset in taking appropriate management actions. Aerobic scope (AS) in fish has been linked to various fitness-related parameters, and may be valuable in determining dispersal potential of aquatic invasive species in novel environments. Round goby, Neogobius melanostomus, one of the most wide-ranging invasive fish species in Europe and North America, currently thrives in brackish and fresh water, but its ability to survive in high salinity waters is unknown to date. We show that AS in round goby is reduced by 30% and blood plasma osmolality increased (indicating reduced capacity for osmoregulation) at salinities approaching oceanic conditions, following slow ramping (5 PSU per week) and subsequent long-term acclimation to salinities ranging between 0 and 30 PSU (8 days at final treatment salinities before blood plasma osmolality measurements, 12–20 additional days before respirometry). Survival was also reduced at the highest salinities yet a significant proportion (61%) of the fish survived at 30 PSU. Reduced physiological performance at the highest salinities may affect growth and competitive ability under oceanic conditions, but to what extent reduced AS and osmoregulatory capacity will slow the current 30 km year-1 rate of advance of the species through the steep salinity gradient from the brackish Baltic Sea and into the oceanic North Sea remains speculative. An unintended natural experiment is in progress to test whether the rate of advance slows down. At the current rate of advance the species will reach the oceanic North Sea by 2018/2019, therefore time for taking preventative action is short.

Atlantic cod ( Gadus morhua L.) exhibits polymorphic hemoglobin variants with the HbI locus showing a strong North-South geographic cline in frequency distribution of three main types (1/1, 1/2 and 2/2). This may indicate selective advantages of the different HbI types under various temperature regimes. Despite this only one study has directly examined the temperature preference of the two homozygous types, HbI-1/1 and HbI-2/2, whereas the preference of the heterozygote (HbI-1/2) has never previously been addressed. By exposing fish to a 4–19 °C temperature gradient in an annular preference chamber we recorded the preferred temperature of wild juvenile G. morhua of all three main Hbl types originating from an area where they co-exist. HbI-2/2 G. morhua preferred significantly cooler water (8.9 ± 0.2 °C) compared to the HbI-1/1 group (11 ± 0.6 °C), this difference, however, not being as distinct as previously reported. There was pronounced inter-individual variation in the temperature preference of the HbI-1/2 G. morhua ranging between 6.7 and 13.8 °C, and their overall preference (10.5 ± 0.9 °C) did not differ significantly from either of the homozygous HbI types. Notably, the mean range of utilized temperature (temperature span between 1st and 3rd quartile) was very similar between all 3 Hbl types with 3.2–3.5 °C. Considering the complexity of a trait like temperature preference, there are clearly many other factors besides HbI type that influence the thermal biology of cod, and therefore we also investigated possible associations between genotype and temperature preference for 12 variable candidate gene single nucleotide polymorphisms (SNPs) a priori expected to be related to growth and reproduction. There were, however, no significant correlations between temperature preference and any of the candidate gene SNPs indicating that none of these polymorphisms strongly associates with thermal behavior. Considering however the high-throughput genotyping methods becoming increasingly accessible there is great potential for association studies involving many more genetic markers to identify additional genetic polymorphisms that are important for temperature preference in G. morhua . In conclusion, we support the notion of a ‘warm’ (HbI-2/2) and a ‘cold’ (HbI-1/1) Hb type, although we suggest the difference to be more subtle than previously reported. Furthermore HbI-1/2 G. morhua shows rather inconsistent thermoregulatory behavior. To obtain a more definitive picture of the extent to which thermal niches are realized under natural conditions field observations in areas where the 3 HbI types co-exist should be performed.

The current study investigated how the gastric evacuation rate (GER) was affected after surgically introducing dummies of a blood flow biotelemetry system into the abdominal cavity of Atlantic cod, Gadus morhua. Gastric evacuation experiments were performed two and 10 days postsurgery on surgically implanted and control G. morhua force-fed sandeel, Ammodytes tobianus. The results were compared with previously obtained estimates from unstressed conspecifics voluntarily feeding on a similar diet. After two days, GER was significantly lower in the group of fish with the dummy implants compared with the control group, but following 10 days of recovery no significant difference was seen between the two groups. The difference between implanted and control fish observed two days postsurgery may have resulted either from surgery, postsurgical stress and/or the presence of the implant. The conclusion is that 10 days of postsurgical recovery will stabilize GER in G. morhua, thus indicating that at this point the implant per se did not affect GER. Both the fish with surgical implants and controls in this study evacuated their stomachs much slower and with much higher interindividual variation compared with G. morhua feeding voluntarily on similar prey items. The lower GER and higher interindividual variation for force-fed fish indicate that handling, anaesthetization and force-feeding impair GER and that individual fish respond differently to the suppressing effects.

The commercially and ecologically valuable sandeel (Ammodytes ssp.) make distinct vertical shifts between an inactive stage, during which they seek refuge in the sand, and a pelagic schooling stage, during which they forage. This characteristic discontinuous foraging pattern constitutes a challenge to fishery biologists and has consequences for a wide range of predators ranging from birds and mammals to commercially important species. However, experimental studies that shed light on the primary drivers of foraging activity in fish are rare. In the present study, whole schools of sandeel (A. tobianus) were caught in August in east Denmark (65°02′30N; 12°37′00E) and kept in large tanks in the laboratory. It was found that the amount of food ingested and memory of past days feeding history are primary drivers of foraging activity at the level of the entire school, whereas external factors such as prey concentration and temperature are merely secondary drivers.

Lesser sandeel (Ammodytes tobianus) is abundant in near-shore areas where it is a key prey. It exhibits the behaviour of alternating between swimming in schools and lying buried in the sediment. We first determined the species’ standard metabolic rate (SMR), critical partial pressure of oxygen $$(P_{{\rm O}_{2{\rm crit}}})$$ and maximal oxygen uptake $$(M_{{\rm O}_{2{\rm max}}}).$$ The sandeel were then exposed to an acute stepwise decline in water oxygen pressure (18.4, 13.8, 9.8, 7.5, 5.8, 4.0, and 3.1 kPa $$P_{{\rm O}_{2}}$$ ). Swimming speed and routine- and post-experimental blood lactate levels were measured, in addition to levels associated with strenuous exercise. The SMR was 69.0 ± 8.4 mg O2 kg−1 h−1 and the $$M_{{\rm O}_{2{\rm max}}}$$ about seven times as high. The $$P_{{\rm O}_{2{\rm crit}}}$$ was found to be 4.1 kPa. A rapid decrease (within 1 h) in $$P_{{\rm O}_{2}}$$ from 18.4 to 3.1 kPa had no significant effect on routine swimming speed (0.9 ± 0.06 bl s−1), but steady levels at the lowest $$P_{{\rm O}_{2}}$$ (3.1 kPa) gradually reduced the swimming speed by 95% after 40 min. The routine blood lactate levels were 2.2 ± 0.6 mmol l−1, while the levels in the strenuously exercised groups were significantly higher with 5.4 ± 1.6 and 5.8 ± 1.3 mmol l−1. The highest levels were observed in post-experimental fish with 7.5 ± 2.7 mmol l−1. We argue that, as sandeel showed no decrease in swimming speed (to offset stress) nor an increased speed to escape the hypoxia, the fish either rely on a low SMR and being a reasonable strong oxygen regulator $$(\hbox{low}\;P_{{\rm O}_{2{\rm crit}}})$$ as a mean to cope when exposed to acute hypoxia, or that the hypoxia simply developed too fast for the fish to decide on an appropriate strategy. Not showing a behavioural response may in the present case be maladaptive, as the consequence was major physiological stress which the fish however appears tolerant towards. The high routine blood lactate levels suggest that anaerobic metabolism is associated with swimming in sandeel, which may be related to the specific lifestyle of the fish where they regularly bury in the sediment.

The round goby, Neogobius melanostomus (Pallas, 1814), is one of the most invasive non-indigenous species in the Baltic Sea. It dominatescoastal fisheries in some localities and is frequently found in offshore pelagic catches. This paper identifies management issues and suggestsactions to be considered for post-invasion management. Priority should be given to the establishment of a coordinated pan-Baltic monitoringprogramme and associated data storage and exchange, as well as the compilation of landing statistics of the round goby in commercial andrecreational fisheries. While eradication is unrealistic, population control that leads to minimising the risk of transfer to yet uncolonised areasin the Baltic Sea and adjacent waterbodies is feasible. This should comprise the requirement that the species be landed in commercial fisherybycatch, the management of ships’ ballast water and sediments, and hull fouling of inland and sea-going vessels, including recreational boats.Extensive involvement of stakeholders is crucial at all phases of the management process.

Haemoglobin polymorphism in cod (Gadus morhua L) has been investigated throughout the last 50 years. Field studies have shed light on the geographic distribution of the two common alleles (HbI1 and HbI2), and laboratory studies have shown effects of genotype on physiological traits such as growth, reproduction and hypoxia tolerance. The geographic distribution of alleles shows a correlation with temperature, with increasing frequency of HbI1 in warmer areas. This is likely due to temperature-related differences in oxygen affinity of the three genotypes. We provide a general ecological introduction to cod haemoglobin polymorphism and a detailed discussion of physiological studies, particularly laboratory growth studies. Although differences in oxygen uptake are almost certainly a contributory mechanism to observed differences in traits such as growth rate, many other environmental, behavioural and social factors may also contribute, making it difficult to quantify the effect of HbI either experimentally or in the field.

Some Atlantic cod in the Bornholm Basin undertake vertical foraging migrations into severely hypoxic bottom water. Hypoxic conditions can reduce the postprandial increase in gastrointestinal blood flow (GBF). This could subsequently postpone or reduce the postprandial increase in oxygen consumption (MO(2)), i.e. the SDA, leading to a disturbed digestion. Additionally, a restricted oxygen uptake could result in an oxygen debt that needs to be compensated for upon return to normoxic waters and this may also affect the ability to process the food. Long-term cardio-respiratory measurements were made on fed G. morhua in order to understand how the cardio-respiratory system of feeding fish respond to a period of hypoxia and a subsequent return to normoxia. These were exposed to 35% water oxygen saturation for 90 minutes, equivalent to the time and oxygen level cod voluntarily endure when searching for food in the Bornholm Basin. We found that i) gastric and intestinal blood flows, cardiac output and MO(2) increased after feeding, ii) gastric and intestinal blood flows were spared in hypoxia, and iii) there were no indications of an oxygen debt at the end of the hypoxic period. The magnitude and time course of the measured variables are similar to values obtained from fish not exposed to the hypoxic period. In conclusion, when cod in the field search for and ingest prey under moderate hypoxic conditions they appear to stay within safe limits of oxygen availability as we saw no indications of an oxygen debt, or negative influence on digestive capacity, when simulating field observations.

Starting in 1980s, the inner Danish waters have yearly been exposed to seasonal oxygen deficiency (hypoxia). Through spatial–temporal interpolation of monitoring data (1998–2005), we investigated oxygen deficiency impacts on suitable burying habitats for lesser sandeel ( Ammodytes tobianus ). Furthermore, the consequences of a predicted 4 °C temperature increase within this century were investigated. Maps of bottom oxygen deficiency (oxygen saturation ≤ Scrit of sandeel) were overlaid on maps of sediment composition. Throughout the study period (1998–2005), about 8% of the suitable sediments were affected by oxygen deficiency during an average year and 23% in the most severe year. Regional differences underlay the interannual variations. The extent of oxygen deficiency in enclosed regions varied from 20% to 33% of the suitable seabed being affected, whereas in open-water regions oxygen deficiency problems were limited during average years. However, large areas of the open-water seabed experienced oxygen deficiency during severe years. In such years, under a 4.0 °C temperature increase scenario, the extent of oxygen deficiency on open-water suitable patches was predicted to increase from 25% to about 40%.

The completed fruit fly genome was found to contain up to 15 putative UDP-N-acetyl-alpha-d-galactosamine:polypeptide N-acetylgalactosaminyltransferase (GalNAc-transferase) genes. Phylogenetic analysis of the putative catalytic domains of the large GalNAc-transferase enzyme families of Drosophila melanogaster (13 available), Caenorhabditis elegans (9 genes), and mammals (12 genes) indicated that distinct subfamilies of orthologous genes are conserved in each species. In support of this hypothesis, we provide evidence that distinctive functional properties of Drosophila and human GalNAc-transferase isoforms were exhibited by evolutionarily conserved members of two subfamilies (dGalNAc-T1 (l(2)35Aa) and GalNAc-T11; dGalNAc-T2 (CG6394) and GalNAc-T7). dGalNAc-T1 and novel human GalNAc-T11 were shown to encode functional GalNAc-transferases with the same polypeptide acceptor substrate specificity, and dGalNAc-T2 was shown to encode a GalNAc-transferase with similar GalNAc glycopeptide substrate specificity as GalNAc-T7. Previous data suggested that the putative GalNAc-transferase encoded by l(2)35Aa had a lethal phenotype (Flores, C., and Engels, W. (1999) Proc. Natl. Acad. Sci. U. S. A. 96, 2964-2969), and this was substantiated by sequencing of three lethal alleles l(2)35Aa(HG8), l(2)35Aa(SF12), and l(2)35Aa(SF32). The finding that subfamilies of GalNAc-transferases with distinct catalytic functions are evolutionarily conserved stresses that GalNAc-transferase isoforms may serve unique biological functions rather than providing functional redundancy, and this is further supported by the lethal phenotype of l(2)35Aa.

The completed fruit fly genome was found to contain up to 15 putative UDP-N-acetyl-α-d-galactosamine:polypeptideN-acetylgalactosaminyltransferase (GalNAc-transferase) genes. Phylogenetic analysis of the putative catalytic domains of the large GalNAc-transferase enzyme families of Drosophila melanogaster (13 available), Caenorhabditis elegans(9 genes), and mammals (12 genes) indicated that distinct subfamilies of orthologous genes are conserved in each species. In support of this hypothesis, we provide evidence that distinctive functional properties of Drosophila and human GalNAc-transferase isoforms were exhibited by evolutionarily conserved members of two subfamilies (dGalNAc-T1 (l(2)35Aa) and GalNAc-T11;dGalNAc-T2 (CG6394) and GalNAc-T7).dGalNAc-T1 and novel human GalNAc-T11 were shown to encode functional GalNAc-transferases with the same polypeptide acceptor substrate specificity, and dGalNAc-T2 was shown to encode a GalNAc-transferase with similar GalNAc glycopeptide substrate specificity as GalNAc-T7. Previous data suggested that the putative GalNAc-transferase encoded by l(2)35Aa had a lethal phenotype (Flores, C., and Engels, W. (1999) Proc. Natl. Acad. Sci. U. S. A. 96, 2964–2969), and this was substantiated by sequencing of three lethal alleles l(2)35Aa HG8,l(2)35Aa SF12, and l(2)35Aa SF32. The finding that subfamilies of GalNAc-transferases with distinct catalytic functions are evolutionarily conserved stresses that GalNAc-transferase isoforms may serve unique biological functions rather than providing functional redundancy, and this is further supported by the lethal phenotype of l(2)35Aa.

Information from data storage tags (DSTs) is conventionally used to infer movement patterns or reveal characteristics (e.g. temperature or salinity) of the environment surrounding tagged fish. Here we link data derived from DSTs with the reproductive state of tagged fish. Indi- vidual vertical activity of adult male and female Atlantic cod Gadus morhua L. in the Bornholm Basin was derived from DST measures and related to the individual histologically determined reproductive phase. Spawning migrations were identified by movements towards deeper and more saline waters. No difference was observed between sexes in the timing of the onset of migra- tion and the duration of migration from feeding grounds to the spawning area. While there was no significant difference in duration of the spawning period between females and males, the histo- logical characteristics suggest that females finish spawning before males. Irrespective of gender, vertical swimming activity was most pronounced during spawning, with descents towards the bot- tom dominating the movements. During spawning, males stayed significantly deeper than females. In conclusion, the present results suggest that initiation of spawning migration and dura- tion of the spawning period differ between sexes, as does the level of activity during spawning events. Not all individuals followed the general pattern; a considerable number of individuals were found to spawn in shallow water in the Arkona Basin, and juvenile fish undertook the migration without spawning.