Proto-cooperation: group hunting sailfish improve hunting success by alternating attacks on grouping prey.

We present evidence of a novel form of group hunting. Individual sailfish (Istiophorus platypterus) alternate attacks with other group members on their schooling prey (Sardinella aurita). While only 24% of attacks result in prey capture, multiple prey are injured in 95% of attacks, resulting in an increase of injured fish in the school with the number of attacks. How quickly prey are captured is positively correlated with the level of injury of the school, suggesting that hunters can benefit from other conspecifics' attacks on the prey. To explore this, we built a mathematical model capturing the dynamics of the hunt. We show that group hunting provides major efficiency gains (prey caught per unit time) for individuals in groups of up to 70 members. We also demonstrate that a free riding strategy, where some individuals wait until the prey are sufficiently injured before attacking, is only beneficial if the cost of attacking is high, and only then when waiting times are short. Our findings provide evidence that cooperative benefits can be realized through the facilitative effects of individuals' hunting actions without spatial coordination of attacks. Such 'proto-cooperation' may be the pre-cursor to more complex group-hunting strategies.

Fast growers sprint slower: effects of food deprivation and re-feeding on sprint swimming performance in individual juvenile European sea bass.

While many ectothermic species can withstand prolonged fasting without mortality, food deprivation may have sublethal effects of ecological importance, including reductions in locomotor ability. Little is known about how such changes in performance in individual animals are related to either mass loss during food deprivation or growth rate during re-feeding. This study followed changes in the maximum sprint swimming performance of individual European sea bass, Dicentrarchus labrax, throughout 45 days of food deprivation and 30 days of re-feeding. Maximum sprint speed did not show a significant decline until 45 days of food deprivation. Among individuals, the reduction in sprinting speed at this time was not related to mass loss. After 30 days of re-feeding, mean sprinting speed had recovered to match that of control fish. Among individuals, however, maximum sprinting speed was negatively correlated with growth rate after the resumption of feeding. This suggests that the rapid compensatory growth that occurs during re-feeding after a prolonged fast carries a physiological cost in terms of reduced sprinting capacity, the extent of which shows continuous variation among individuals in relation to growth rate. The long-term repeatability of maximum sprint speed was low when fish were fasted or fed a maintenance ration, but was high among control fish fed to satiation. Fish that had been previously food deprived continued to show low repeatability in sprinting ability even after the initiation of ad libitum feeding, probably stemming from variation in compensatory growth among individuals and its associated negative effects on sprinting ability. Together, these results suggest that food limitation can disrupt hierarchies of maximum sprint performance within populations. In the wild, the cumulative effects on locomotor capacity of fasting and re-feeding could lead to variable survival among individuals with different growth trajectories following a period of food deprivation.

Hidden impacts of ocean warming and acidification on biological responses of marine animals revealed through meta-analysis.

Conflicting results remain on the impacts of climate change on marine organisms, hindering our capacity to predict the future state of marine ecosystems. To account for species-specific responses and for the ambiguous relation of most metrics to fitness, we develop a meta-analytical approach based on the deviation of responses from reference values (absolute change) to complement meta-analyses of directional (relative) changes in responses. Using this approach, we evaluate responses of fish and invertebrates to warming and acidification. We find that climate drivers induce directional changes in calcification, survival, and metabolism, and significant deviations in twice as many biological responses, including physiology, reproduction, behavior, and development. Widespread deviations of responses are detected even under moderate intensity levels of warming and acidification, while directional changes are mostly limited to more severe intensity levels. Because such deviations may result in ecological shifts impacting ecosystem structures and processes, our results suggest that climate change will likely have stronger impacts than those previously predicted based on directional changes alone.

Aerobic metabolic scope mapping of an invasive fish species with global warming.

Climate change will exacerbate the negative effects associated with the introduction of non-indigenous species in marine ecosystems. Predicting the spread of invasive species in relation to environmental warming is therefore a fundamental task in ecology and conservation. The Baltic Sea is currently threatened by several local stressors and the highest increase in sea surface temperature of the world's large marine ecosystems. These new thermal conditions can further favour the spreading of the invasive round goby (Neogobius melanostomus), a fish of Ponto-Caspian origin, currently well established in the southern and central parts of the Baltic Sea. This study aims to assess the thermal habitat suitability of the round goby in the Baltic Sea considering the past and future conditions. The study combines sightings records with known physiological models of aerobic performance and sea surface temperatures. Physiological models read these temperatures, at sighting times and locations, to determine their effects on the aerobic metabolic scope (AMS) of the fish, a measure of its energetic potential in relation to environmental conditions. The geographical mapping of the AMS was used to describe the changes in habitat suitability during the past 3 decades and for climatic predictions (until 2100) showing that the favourable thermal habitat in the Baltic Sea has increased during the past 32 years and will continue to do so in all the applied climate model predictions. Particularly, the predicted new thermal conditions do not cause any reduction in the AMS of round goby populations, while the wintertime cold ranges are likely expected to preserve substantial areas from invasion. The results of this research can guide future monitoring programs increasing the chance to detect this invader in novel areas.

Ocean acidification and seasonal temperature extremes combine to impair the thermal physiology of a sub-Antarctic fish.

To predict the potential impacts of climate change on marine organisms, it is critical to understand how multiple stressors constrain the physiology and distribution of species. We evaluated the effects of seasonal changes in seawater temperature and near-future ocean acidification (OA) on organismal and sub-organismal traits associated with the thermal performance of Eleginops maclovinus, a sub-Antarctic notothenioid species with economic importance to sport and artisanal fisheries in southern South America. Juveniles were exposed to mean winter and summer sea surface temperatures (4 and 10 °C) at present-day and near-future pCO2 levels (~500 and 1800 µatm). After a month, the Critical Thermal maximum and minimum (CTmax, CTmin) of fish were measured using the Critical Thermal Methodology and the aerobic scope of fish was measured based on the difference between their maximal and standard rates determined from intermittent flow respirometry. Lipid peroxidation and the antioxidant capacity were also quantified to estimate the oxidative damage potentially caused to gill and liver tissue. Although CTmax and CTmin were higher in individuals acclimated to summer versus winter temperatures, the increase in CTmax was minimal in juveniles exposed to the near-future compared to present-day pCO2 levels (there was a significant interaction between temperature and pCO2 on CTmax). The reduction in the thermal tolerance range under summer temperatures and near-future OA conditions was associated with a reduction in the aerobic scope observed at the elevated pCO2 level. Moreover, an oxidative stress condition was detected in the gill and liver tissues. Thus, chronic exposure to OA and the current summer temperatures pose limits to the thermal performance of juvenile E. maclovinus at the organismal and sub-organismal levels, making this species vulnerable to projected climate-driven warming.

Aerobic capacity influences the spatial position of individuals within fish schools.

The schooling behaviour of fish is of great biological importance, playing a crucial role in the foraging and predator avoidance of numerous species. The extent to which physiological performance traits affect the spatial positioning of individual fish within schools is completely unknown. Schools of juvenile mullet Liza aurata were filmed at three swim speeds in a swim tunnel, with one focal fish from each school then also measured for standard metabolic rate (SMR), maximal metabolic rate (MMR), aerobic scope (AS) and maximum aerobic swim speed. At faster speeds, fish with lower MMR and AS swam near the rear of schools. These trailing fish required fewer tail beats to swim at the same speed as individuals at the front of schools, indicating that posterior positions provide hydrodynamic benefits that reduce swimming costs. Conversely, fish with high aerobic capacity can withstand increased drag at the leading edge of schools, where they could maximize food intake while possibly retaining sufficient AS for other physiological functions. SMR was never related to position, suggesting that high maintenance costs do not necessarily motivate individuals to occupy frontal positions. In the wild, shifting of individuals to optimal spatial positions during changing conditions could influence structure or movement of entire schools.

Conservation physiology of marine fishes: advancing the predictive capacity of models.

At the end of May, 17 scientists involved in an EU COST Action on Conservation Physiology of Marine Fishes met in Oristano, Sardinia, to discuss how physiology can be better used in modelling tools to aid in management of marine ecosystems. Current modelling approaches incorporate physiology to different extents, ranging from no explicit consideration to detailed physiological mechanisms, and across scales from a single fish to global fishery resources. Biologists from different sub-disciplines are collaborating to rise to the challenge of projecting future changes in distribution and productivity, assessing risks for local populations, or predicting and mitigating the spread of invasive species.

Behavioural and kinematic components of the fast-start escape response in fish: individual variation and temporal repeatability.

Inter-individual variation in physiological performance traits, which is stable over time, can be of potential ecological and evolutionary significance. The fish escape response is interesting in this regard because it is a performance trait for which inter-individual variation may determine individual survival. The temporal stability of such variation is, however, largely unexplored. We quantified individual variation of various components of the escape response in a population of European sea bass (Dicentrarchus labrax), considering both non-locomotor (responsiveness and latency) and locomotor (speed, acceleration, turning rate, turning angle and distance travelled in a fixed time, D(esc)) variables. We assessed whether variation in performance was temporally stable and we searched for any trade-offs among the components of the response that might explain why the variation persisted in the population. The coefficient of variation was high for all components, from 23% for turning rate to 41% for D(esc), highlighting the non-stereotypic nature of the response. Individual performance for all variables was significantly repeatable over five sequential responses at 30 min intervals, and also repeatable after a 30 day interval for most of the components. This indicates that the variation is intrinsic to the individuals, but there was no evidence for trade-offs amongst the components of the response, suggesting that, if trade-offs exist, they must be against other ecologically important behavioural or performance traits.

Fuel, fasting, fear: routine metabolic rate and food deprivation exert synergistic effects on risk-taking in individual juvenile European sea bass.

1. Individuals of the same species often exhibit consistent differences in metabolic rate, but the effects of such differences on ecologically important behaviours remain largely unknown. In particular, it is unclear whether there is a cause-and-effect relationship between metabolic rate and the tendency to take risks while foraging. Individuals with higher metabolic rates may need to take greater risks while foraging to obtain the additional food required to satisfy their energy requirements. Such a relationship could be exacerbated by food deprivation if a higher metabolic demand also causes greater mass loss and hunger. 2. We investigated relationships among metabolic rate, risk-taking and tolerance of food deprivation in juvenile European sea bass. Individual fish were tested for risk-taking behaviours following a simulated predator attack, both before and after a 7-day period of food deprivation. The results were then related to their routine metabolic rate (RMR), which was measured throughout the period of food deprivation. 3. The amount of risk displayed by individual fish before food deprivation showed no relationship with RMR. After food deprivation, however, the amount of risk among individuals was positively correlated with RMR. In general, most fish showed an increase in risk-taking after food deprivation, and the magnitude of the increase in risk-taking was correlated with the rate of individual mass loss during food deprivation, which was itself strongly correlated with RMR. 4. The observation that RMR was related to risk-taking behaviour after food deprivation, but not before, suggests that although RMR can influence risk-taking, the strength of the relationship is flexible and context dependent. The effects of RMR on risk-taking may be subtle or non-existent in regularly feeding animals, but may lead to variability in risk-taking among individuals when food is scarce or supply is unpredictable. This synergistic relationship between RMR and food deprivation could lead to an increased likelihood of being predated for individuals with a relatively high intrinsic energy demand during times when food is scarce.

The effects of hypoxia on aerobic metabolism in oil-contaminated sea bass (Dicentrarchus labrax).

Hypoxia and petrogenic hydrocarbon contamination are two anthropogenic stressors that coexist in coastal environments. Although studies have estimated the impact of each stressor separately, few investigations have assessed the effects of these stressors in interaction. We therefore investigated the impact of these combined stressors on sea bass, (Dicentrarchus labrax) physiology. After experimental contamination with physically dispersed oil, fish were exposed to hypoxia or normoxia, and active/standard metabolic rates (AMR and SMR, respectively), and metabolic scope (MS) were estimated. At the protocol's end, the uptake of polycyclic aromatic hydrocarbons (PAHs) was estimated by evaluating relative concentrations of bile metabolites. In terms of bile metabolites, our results validated the uptake of PAHs by contaminated fish in our experimental settings, and further suggest that the hypoxic period after contamination does not reduce or increase compound metabolization processes. Our data showed significant effects of hypoxia on all metabolic rates: a significant drastic AMR reduction and significant SMR diminution led to decreased MS. We also found that oil contamination significantly impacted AMR and MS, but not SMR. These results suggested that when evaluated separately, hypoxia or oil affect the metabolic rate of sea bass. On the other hand, when evaluated in combination, no cumulative effects were observed, since fish exposed to both stressors did not show a stronger impact on metabolism than fish exposed to hypoxia alone. This suggests that oil impacts fish metabolism when fish occupy normoxic waters, and that oil does not magnify hypoxia-induced effects on fish metabolism.

Combined effect of pCO2 and temperature levels on the thermal niche in the early benthic ontogeny of a keystone species.

We evaluated the effects of projected, near future ocean acidification (OA) and extreme events of temperature (warming or cooling) on the thermal tolerance of Concholepas concholepas, a coastal benthic keystone species. Three separate trials of an experiment were conducted by exposing juvenile C. concholepas for 1 month to one of two contrasting pCO2 levels (~500 and ~1200 µatm). In addition, each pCO2 level was combined with one of four temperature treatments. The control was 15 °C, whilst the other temperatures were 10 °C (Trial 1), 20 °C (Trial 2) and 25 °C (Trial 3). At the end of each trial, we assessed Critical Thermal maximum (CTmax) and minimum (CTmin) via self-righting success, calculated partial thermal tolerance polygons, measured somatic growth, determined transcription of Heat Shock Proteins 70 (HSP70) and measured oxygen consumption rates. Regardless of pCO2 level, HSP70 transcript levels were significantly higher in juveniles after exposure to extreme temperatures (10 °C and 25 °C) indicating physiological stress. Oxygen consumption rates increased with increasing temperature from 10 °C to 20 °C though showed a decrease at 25 °C. This rate was not affected by pCO2 or the interaction between temperature and pCO2. Juveniles exposed to present-day and near future pCO2 levels at 20 °C showed similar thermal tolerance polygonal areas; whilst changes in both CTmin and CTmax at 25 °C and 10 °C caused narrower and broader areas, respectively. Temperature affected growth, oxygen consumption and HSP70 transcription in small juvenile C. concholepas. Exposure to elevated pCO2 did not affect thermal tolerance, growth or oxygen consumption at temperatures within the thermal range normally experienced by this species in northern Chile (15-20 °C). At elevated pCO2 conditions, however, exposure to warmer (25 °C) or colder (10 °C) temperatures reduced or increased the thermal area, respectively. This study demonstrates the importance of examining the thermal-tolerance edges to better understand how OA and temperature will combine to physiologically challenge inter-tidal organisms.

The effect of hypoxia and hydrocarbons on the anti-predator performance of European sea bass (Dicentrarchus labrax).

Hydrocarbons contamination and hypoxia are two stressors that can coexist in coastal ecosystems. At present, few studies evaluated the combined impact of these stressors on fish physiology and behavior. Here, we tested the effect of the combination of hypoxia and petrogenic hydrocarbons on the anti-predator locomotor performance of fish. Specifically, two groups of European sea bass (Dicentrarchus labrax) were exposed to clean water (Ctrl) or oil-contaminated water (Oil). Subsequently, fish of both groups were placed in normoxic (norx) or hypoxic (hyp) experimental tanks (i.e. four groups of fish were formed: Ctrl norx, Ctrl hyp, Oil norx, Oil hyp). In these tanks, escape response was elicited by a mechano-acoustic stimulus and recorded with a high speed camera. Several variables were analyzed: escape response duration, responsiveness (percentage of fish responding to the stimulation), latency (time taken by the fish to initiate a response), directionality (defined as away or toward the stimulus), distance-time variables (such as speed and acceleration), maneuverability variables (such as turning rate), escape trajectory (angle of flight) and distancing of the fish from the stimulus. Results revealed (i) effects of stressors (Ctrl hyp, Oil norx and Oil hyp) on the directionality; (ii) effects of Oil norx and Oil hyp on maneuverability and (iii) effects of Oil hyp on distancing. These results suggest that individual stressors could alter the escape response of fish and that their combination could strengthen these effects. Such an impact could decrease the probability of prey escape success. By investigating the effects of hydrocarbons (and the interaction with hypoxia) on the anti-predator behavior of fish, this work increases our understanding of the biological impact of oil spill. Additionally, the results of this study are of interest for oil spill impact evaluation and also for developing new ecotoxicological tools of ecological significance.

Ocean warming and acidification pose synergistic limits to the thermal niche of an economically important echinoderm.

To make robust projectios of the impacts of climate change, it is critical to understand how abiotic factors may interact to constrain the distribution and productivity of marine flora and fauna. We evaluated the effects of projected end of the century ocean acidification (OA) and warming (OW) on the thermal tolerance of an important living marine resource, the sea urchin Loxechinus albus, a benthic shallow water coastal herbivore inhabiting part of the Pacific coast of South America. After exposing young juveniles for a 1-month period to contrasting pCO2 (~500 and 1400 µatm) and temperature (~15 °C and 20 °C) levels, critical thermal maximum (CTmax) and minimum (CTmin) as well as thermal tolerance polygons were assessed based on self-righting success as an end point. Transcription of heat shock protein 70 (HSP70), a chaperone protecting cellular proteins from environmental stress, was also measured. Exposure to elevated pCO2 significantly reduced thermal tolerance by increasing CTmin at both experimental temperatures and decreasing CTmax at 20 °C. There was also a strong synergistic effect of OA × OW on HSP70 transcription levels which were 75 times higher than in control conditions. If this species is unable to adapt to elevated pCO2 in the future, the reduction in thermal tolerance and HSP response suggests that near-future warming and OA will disrupt their performance and reduce their distribution with ecological and economic consequences. Given the wider latitudinal range (6 to 56°S) and environmental tolerance of L. albus compared to other members of this region's benthic invertebrate community, OW and OA may cause substantial changes to the coastal fauna along this geographical range.

Avoidance threshold to oil water-soluble fraction by a juvenile marine teleost fish.

When oil spills occur, behavior is the first line of defense for a fish to avoid being contaminated. We determined the avoidance threshold of the European seabass (Dicentrarchus labrax) to the water-soluble fraction (WSF) of oil using a dual-flow choice box. The results showed that a plume of 20%-diluted WSF (total polycyclic aromatic hydrocarbon [PAH] concentration: 8.54 µg L-1 ) triggered a significant avoidance response that was detected within 7.5 min of introducing WSF-contaminated water into the experimental setup. However, the ecological relevance of seabass capacity to detect and avoid WSF remains to be established. In the short term, such a response is indeed liable to reduce seabass contact time with oil-contaminated water and thus preserve their functional integrity. In the long term, however, avoidance may contribute to the displacement of a population into a possibly less auspicious environment, with consequences very similar to those of contaminant exposure, that is, disturbed population dynamics and demography. Environ Toxicol Chem 2018;37:854-859. © 2017 SETAC.

The effect of hypoxia on fish schooling.

Low-oxygen areas are expanding in the oceans as a result of climate change. Work carried out during the past two decades suggests that, in addition to impairing basic physiological functions, hypoxia can also affect fish behaviour. Given that many fish species are known to school, and that schooling is advantageous for their survival, the effect of hypoxia on schooling behaviour may have important ecological consequences. Here, we review the effects of hypoxia on school structure and dynamics, together with the mechanisms that cause an increase in school volume and that ultimately lead to school disruption. Furthermore, the effect of hypoxia generates a number of trade-offs in terms of schooling positions and school structure. Field observations have found that large schools of fish can exacerbate hypoxic conditions, with potential consequences for school structure and size. Therefore, previous models that predict the maximum size attainable by fish schools in relation to oxygen levels are also reviewed. Finally, we suggest that studies on the effect of hypoxia on schooling need to be integrated with those on temperature and ocean acidifications within a framework aimed at increasing our ability to predict the effect of multiple stressors of climate change on fish behaviour.This article is part of the themed issue 'Physiological determinants of social behaviour in animals'.

The role of physiological traits in assortment among and within fish shoals.

Individuals of gregarious species often group with conspecifics to which they are phenotypically similar. This among-group assortment has been studied for body size, sex and relatedness. However, the role of physiological traits has been largely overlooked. Here, we discuss mechanisms by which physiological traits-particularly those related to metabolism and locomotor performance-may result in phenotypic assortment not only among but also within animal groups. At the among-group level, varying combinations of passive assortment, active assortment, phenotypic plasticity and selective mortality may generate phenotypic differences among groups. Even within groups, however, individual variation in energy requirements, aerobic and anaerobic capacity, neurological lateralization and tolerance to environmental stressors are likely to produce differences in the spatial location of individuals or associations between group-mates with specific physiological phenotypes. Owing to the greater availability of empirical research, we focus on groups of fishes (i.e. shoals and schools). Increased knowledge of physiological mechanisms influencing among- and within-group assortment will enhance our understanding of fundamental concepts regarding optimal group size, predator avoidance, group cohesion, information transfer, life-history strategies and the evolutionary effects of group membership. In a broader perspective, predicting animal responses to environmental change will be impossible without a comprehensive understanding of the physiological basis of the formation and functioning of animal social groups.This article is part of the themed issue 'Physiological determinants of social behaviour in animals'.

The Evolution of Lateralization in Group Hunting Sailfish.

Lateralization is widespread throughout the animal kingdom [1-7] and can increase task efficiency via shortening reaction times and saving on neural tissue [8-16]. However, lateralization might be costly because it increases predictability [17-21]. In predator-prey interactions, for example, predators might increase capture success because of specialization in a lateralized attack, but at the cost of increased predictability to their prey, constraining the evolution of lateralization. One unexplored mechanism for evading such costs is group hunting: this would allow individual-level specialization, while still allowing for group-level unpredictability. We investigated this mechanism in group hunting sailfish, Istiophorus platypterus, attacking schooling sardines, Sardinella aurita. During these attacks, sailfish alternate in attacking the prey using their elongated bills to slash or tap the prey [22-24]. This rapid bill movement is either leftward or rightward. Using behavioral observations of identifiable individual sailfish hunting in groups, we provide evidence for individual-level attack lateralization in sailfish. More strongly lateralized individuals had a higher capture success. Further evidence of lateralization comes from morphological analyses of sailfish bills that show strong evidence of one-sided micro-teeth abrasions. Finally, we show that attacks by single sailfish are indeed highly predictable, but predictability rapidly declines with increasing group size because of a lack of population-level lateralization. Our results present a novel benefit of group hunting: by alternating attacks, individual-level attack lateralization can evolve, without the negative consequences of individual-level predictability. More generally, our results suggest that group hunting in predators might provide more suitable conditions for the evolution of strategy diversity compared to solitary life.

Maximum swimming speeds of sailfish and three other large marine predatory fish species based on muscle contraction time and stride length: a myth revisited.

Billfishes are considered to be among the fastest swimmers in the oceans. Previous studies have estimated maximum speed of sailfish and black marlin at around 35 m s-1 but theoretical work on cavitation predicts that such extreme speed is unlikely. Here we investigated maximum speed of sailfish, and three other large marine pelagic predatory fish species, by measuring the twitch contraction time of anaerobic swimming muscle. The highest estimated maximum swimming speeds were found in sailfish (8.3±1.4 m s-1), followed by barracuda (6.2±1.0 m s-1), little tunny (5.6±0.2 m s-1) and dorado (4.0±0.9 m s-1); although size-corrected performance was highest in little tunny and lowest in sailfish. Contrary to previously reported estimates, our results suggest that sailfish are incapable of exceeding swimming speeds of 10-15 m s-1, which corresponds to the speed at which cavitation is predicted to occur, with destructive consequences for fin tissues.

Conservation physiology of marine fishes: state of the art and prospects for policy.

The state of the art of research on the environmental physiology of marine fishes is reviewed from the perspective of how it can contribute to conservation of biodiversity and fishery resources. A major constraint to application of physiological knowledge for conservation of marine fishes is the limited knowledge base; international collaboration is needed to study the environmental physiology of a wider range of species. Multifactorial field and laboratory studies on biomarkers hold promise to relate ecophysiology directly to habitat quality and population status. The 'Fry paradigm' could have broad applications for conservation physiology research if it provides a universal mechanism to link physiological function with ecological performance and population dynamics of fishes, through effects of abiotic conditions on aerobic metabolic scope. The available data indicate, however, that the paradigm is not universal, so further research is required on a wide diversity of species. Fish physiologists should interact closely with researchers developing ecological models, in order to investigate how integrating physiological information improves confidence in projecting effects of global change; for example, with mechanistic models that define habitat suitability based upon potential for aerobic scope or outputs of a dynamic energy budget. One major challenge to upscaling from physiology of individuals to the level of species and communities is incorporating intraspecific variation, which could be a crucial component of species' resilience to global change. Understanding what fishes do in the wild is also a challenge, but techniques of biotelemetry and biologging are providing novel information towards effective conservation. Overall, fish physiologists must strive to render research outputs more applicable to management and decision-making. There are various potential avenues for information flow, in the shorter term directly through biomarker studies and in the longer term by collaborating with modellers and fishery biologists.