Effect of turbidity on habitat preference of juvenile Atlantic cod, Gadus morhua

We examined the effects of turbidity on habitat preference of juvenile Atlantic cod in the laboratory, using a shuttle box where fish could select between two different habitats. In the first experiment, we compared three turbidity levels of kaolin (3, 8 and 21 beam attenuation m⁻¹). In the second experiment, we looked at the effect of turbidity media (kaolin versus algae), after controlling for spectral differences between turbidity media. Although cod preferred an intermediate turbidity of kaolin over low turbidity water, comparisons between low and high turbidity, and intermediate and high turbidity did not significantly influence habitat preference. Algae did not influence habitat preference by cod. Although other studies have found that turbidity affects both foraging and antipredator behaviour of juvenile cod, this study has shown that gradients in turbidity per se do not have a strong effect on their habitat preference.     

Comments on a study of the thermal behaviour of the American eel (Anguilla rostrata) and some statistical suggestions for temperature preference studies

Some suggestions are made concerning methods and statistical evaluation of results in temperature preference studies. The use of regression analysis is advocated rather than graphical methods for evaluating results from acute temperature preference studies and a method of assessing the final preferendum and its confidence interval is demonstrated. This method is applied to a previously published study of the acutely preferred temperature of the American eel (Barila & Stauffer, 1980). The final preferendum and its 95% confidence interval is shown to be 17.4° ± 2.0 °C. In direct contrast to conclusions there drawn, the preferred temperature is shown to depend on the acclimated temperature. Some critical comments are also made concerning other parts of the study.     

Parallel adaptive evolution of Atlantic cod on both sides of the Atlantic Ocean in response to temperature

Despite the enormous economic and ecological importance of marine organisms, the spatial scales of adaptation and biocomplexity remain largely unknown. Yet, the preservation of local stocks that possess adaptive diversity is critical to the long-term maintenance of productive stable fisheries and ecosystems. Here, we document genomic evidence of range-wide adaptive differentiation in a broadcast spawning marine fish, Atlantic cod (Gadus morhua), using a genome survey of single nucleotide polymorphisms. Of 1641 gene-associated polymorphisms examined, 70 (4.2%) tested positive for signatures of selection using a Bayesian approach. We identify a subset of these loci (n = 40) for which allele frequencies show parallel temperature-associated clines (p < 0.001, r² = 0.89) in the eastern and western north Atlantic. Temperature associations were robust to the statistical removal of geographic distance or latitude effects, and contrasted ‘neutral’ loci, which displayed no temperature association. Allele frequencies at temperature-associated loci were significantly correlated, spanned three linkage groups and several were successfully annotated supporting the involvement of multiple independent genes. Our results are consistent with the evolution and/or selective sweep of multiple genes in response to ocean temperature, and support the possibility of a new conservation paradigm for non-model marine organisms based on genomic approaches to resolving functional and adaptive diversity.     

Behavioral responses of Atlantic cod to sea temperature changes

Understanding responses of marine species to temperature variability is essential to predict impacts of future climate change in the oceans. Most ectotherms are expected to adjust their behavior to avoid extreme temperatures and minimize acute changes in body temperature. However, measuring such behavioral plasticity in the wild is challenging. Combining 4 years of telemetry‐derived behavioral data on juvenile and adult (30–80 cm) Atlantic cod (Gadus morhua), and in situ ocean temperature measurements, we found a significant effect of sea temperature on cod depth use and activity level in coastal Skagerrak. During summer, cod were found in deeper waters when sea surface temperature increased. Further, this effect of temperature was stronger on larger cod. Diel vertical migration, which consists in a nighttime rise to shallow feeding habitats, was stronger among smaller cod. As surface temperature increased beyond ~15°C, their vertical migration was limited to deeper waters. In addition to larger diel vertical migrations, smaller cod were more active and travelled larger distances compared to larger specimens. Cold temperatures during winter tended, however, to reduce the magnitude of diel vertical migrations, as well as the activity level and distance moved by those smaller individuals. Our findings suggest that future and ongoing rises in sea surface temperature may increasingly deprive cod in this region from shallow feeding areas during summer, which may be detrimental for local populations of the species.     

The effect of reduced temperature and salinity on the blood physiology of juvenile Atlantic cod

The osmolality and ionic composition of the blood of juvenile Atlantic cod Gadus morhua and their response to conditions of reduced temperature and salinity in summer‐ and winter‐acclimated individuals was investigated. Haematocrit percentage was relatively stable throughout the experimental procedures. Summer‐acclimated juvenile Atlantic cod had higher plasma osmolality than winter‐acclimated fish in ambient conditions. Plasma Na⁺ levels were, however, higher in winter conditions, while Cl⁻ did not vary between seasons. Temperature reduction (12, 9 and 6° C in summer and to 6 and 4° C in winter) induced a significant response in plasma osmolality and Na⁺ levels in summer, but only in Na⁺ levels in winter‐acclimated fish. A pronounced effect was seen in the summer 6° C treatment. Salinity treatments (24, 16 and 8) had a significant effect on almost all the variables in both summer and winter and resulted generally in dilution of ionic and osmotic concentrations of the plasma. This effect was pronounced in the lowest temperature treatments, with the greatest reduction observed in the summer 6° C treatment. This could suggest that winter‐acclimated fish are physiologically adapted to cope with lower seawater temperatures as opposed to summer‐acclimated fish.     

Seasonal growth of Atlantic cod: effects of temperature, feeding and reproduction

Growth of 2659 Atlantic cod Gadus morhua aged 4 to 9 years examined in Placentia Bay, Newfoundland, peaked in most cases in June and was at a minimum in October or November. Water temperature, partial fullness index ( I _P) and gonado‐somatic index ( I _G) explained between 31 and 52% of the monthly variability in growth. Temperature and I _P of capelin Mallotus villosus had significant effects on growth of all age groups and explained most of the variance for ages 6–8 and 4–5 years, respectively. The I _P of large invertebrates (ages 4 to 7 years), sandlance ( Ammodytes sp. age 6 years) and demersal fishes (age 9 years) had age‐specific effects in the model. Overall, amphipods, decapods and echinoderms dominated the Atlantic cod diet in most seasons, but fish consumption by Atlantic cod was high in June and July, particularly on capelin. The rapid increase in somatic mass during June and July occurred despite cold water temperatures ( < 3° C at 50 m) and moderate to high gonado‐somatic index. The findings of this study suggest that when food was not a limiting factor, growth tended to increase even when Atlantic cod occupied colder waters, but when food was limiting, the opposite may have occured.     

Temperature effects on otolith pattern formation in Atlantic cod Gadus morhua

The effect of food intake and temperature on otolith macrostructure and microstructure was examined experimentally in Atlantic cod Gadus morhua. Daily increment formation was validated and otolith accretion rate and optical density quantified using image analysis. Two‐week periods of starvation had no discernable effect on otolith increment width or optical density, despite having negative effects on somatic growth. In contrast, temperature had a strong positive effect on otolith accretion rate and clear effects on optical density with the otolith becoming more translucent at higher temperatures. Somatic growth, otolith accretion and otolith optical density each had a significantly different response curve to temperature. No relationship was detected between individual somatic growth rates and the accretion rate or optical properties of the otolith. The experimental manipulation of temperature‐induced otolith patterns similar to the ‘false ring’ secondary structures sometimes observed in the otoliths of wild fish. The results suggest that otolith pattern arises from a combination of temperature and seasonal effects, but not directly from individual variation in somatic growth.     

Effects of varying temperature and food availability on growth and reproduction in first‐time spawning female Atlantic cod

The somatic growth, sexual maturation and fecundity of individually marked first‐time spawning female Atlantic cod Gadus morhua were examined under different varying temperature and feeding regimes over the months preceding spawning. A negative correlation between somatic and oocyte growth was found, reflecting the changing energy allocation pattern. Nevertheless, the somatic growth of mature individuals was at least as high as those of immature fish over the period of vitellogenesis. Potential fecundity was positively correlated with body size, but neither temperature or feeding regime significantly affected this relationship. Consequently, fish with unlimited feeding opportunity invested more energy into somatic growth during vitellogenesis compared to those held under a restricted ration. This indicated that once Atlantic cod had made the decision to invest in first reproduction, they allocated a certain amount of energy relative to their size into egg production and any surplus was invested into somatic growth. Low temperature led to an arrest in the onset of vitellogenesis and significantly affected the number of females that matured.     

Lethal oxygen levels at different temperatures and the preferred temperature during hypoxia of the Atlantic cod, Gadus morhua L.

The behavioural thermoregulation of Atlantic cod Gadus morhua L. was investigated in a shuttlebox at normoxia and at three levels of hypoxia: 30, 20 and 15% oxygen saturation.
The preferred temperatures at normoxia, 30, 20 and 15% oxygen saturation were 13·9, 13·1. 10·0 and 8·8° C, respectively.
A decrease in metabolism and an increased blood oxygen affinity are among the physiological advantages of selecting a lower temperature during hypoxia. Furthermore the chances of surviving low oxygen saturations are better at low temperatures.
In natural environments, this behaviour may result in habitat shifts of fish living in heterothermal environments with changing oxygen saturations, especially in coastal areas with eutrophication, as for example the Baltic Sea.     

The final thermal preferendum of fishes: Shuttling behavior and acclimation overshoot

The concept of final preferendum is reexamined in light of data concerning acclimation in a cyclic thermal regime. Because of Shuttling behavior to temperatures above and below the mean preferendum, and because of faster acclimation upward than downward, which result in acclimation to a temperature exceeding the mean of the cycle, a fish may finally gravitate to a preferendum which does not equal acclimation temperature. It is suggested that a distinction be made between the crossover-point preferendum, where preference and acclimation are equal, and the ultimate preferendum to which a fish will ultimately gravitate.     

Persistent growth effects of temperature and photoperiod in Atlantic cod Gadus morhua

Short-term environmental manipulations during the early juvenile stage have a large impact on harvesting size of Atlantic cod Gadus morhua nearly 3 years later. A group of juvenile Atlantic cod (initial mass 9·5 g) were reared for 3 months under simulated natural photoperiod or continuous light, and a range of temperatures (7, 10, 13 and 16° C, and a group called T-step, i.e. with temperature reduced successively from 16 to 13 and 10° C). After termination of the laboratory trial, the fish were moved to sea pens and reared at ambient conditions for 30 months before harvesting in June 2006. Observed growth gain from the 3 month laboratory trial was still persistent following the 30 months of sea-pen ongrowing. The T-step group displayed 15, 13, 1 and 10% superior mass gain respectively than the groups initially at 7, 10, 13 and 16° C at harvest in June 2006. Similarly, rearing under continuous light during the initial 3 month period during the early juvenile stage resulted in 1–9% larger size at harvesting compared to fish reared at simulated natural photoperiod. Gonado-somatic and hepato-somatic indices were similar in all groups. Contribution to the understanding of the mechanism behind size variation in adult fish can have wide range applications for Atlantic cod fisheries and aquaculture.     

Hypoxia tolerance in Atlantic cod

Oxygen saturation levels that killed 50 and 5% of cod Gadus morhua over 96 h averaged 21·2 and 27·7%, respectively. No fish survived at 10% saturation and only a few survived at 16% saturation, whereas no mortality occurred at 34 and 40% oxygen saturation. Since metabolic rate and oxygen consumption increase with increasing temperature, we hypothesized that cod would be less tolerant to hypoxic conditions at 6 than at 2° C. However, temperature (2 and 6° C) had no measurable impact on cod survival. Small (mean & S.D.; 45·2 ± 4·2 cm) and large (57·5 ± 3·8 cm) cod had the same tolerance to hypoxia. At the end of the experiments, hypoxia had a significant effect on blood haematocrit, mean cellular haemoglobin content, liver lactate, plasma glucose and plasma lactate, but accounted for only a small fraction (< 10%) of the variation, except for plasma lactate which exhibited a strong response with concentrations increasing progressively with decreasing levels of oxygen saturation. Temperature had a significant effect on most variates in normoxia and hypoxia. Variates also affected by oxygen level showed significant interactions between oxygen and size or temperature effects. However, these interactions accounted for only a small proportion of the variation. Physiological parameters indicated that extending the duration of our tests beyond 96 h would not have changed our estimates of the lethal thresholds. Hypoxic conditions are a permanent feature of the deep waters of the Gulf of St Lawrence. This study shows that a significant portion of the benthic habitats in the Gulf are uninhabitable for cod which would be expected to avoid waters below 28% oxygen saturation.     

Influence of acclimation temperature and season on acute temperature preference of adult mountain whitefish,Prosopium williamsoni

Synopsis Temperatures preferred by four groups of adult mountain whitefish, Prosopium williamsoni, collected in October before spawning, in December after spawning, in late winter, and in spring were determined in the laboratory in a horizontal gradient. Acute temperature preference based on fish tested soon after capture and final preferendum estimates were 12.8 and 17.7°C, respectively (pre-spawning), 9.6 and 11.9°C (post-spawning), 10.8 and 9.9°C (winter), and 16.4 and 16.3°C (spring). Seasonal influence on temperature preference was evident on the basis of differences in final preferenda, covariance analysis of responses of laboratory-acclimated fish, and temperature preference of fish held at ambient river temperatures. Fish of the post-spawning and winter groups preferred lower temperatures than did those of pre-spawning and spring groups. Temperatures preferred by pre-spawning fish were too high for embryo survival. Caution is necessary in predicting thermal preference on the basis of a sample collected at one time of year or stage of sexual development.     

Stock-specific changes in growth rates, food conversion efficiencies, and energy allocation in response to temperature change in juvenile Atlantic cod

Under common environments, populations of laboratory reared (Grand Banks, GB and Gulf of Maine, GOM) and wild caught (Fortune Bay, FB and Bonavista Bay, BB; Newfoundland) juvenile cod Gadus morhua responded similarly to temperature change in specific growth rates, food conversion efficiencies, condition factors, liver water content, and muscle water content. However, GOM cod had higher condition factors, and showed differences from GB cod in phenotypic plasticity of hepatosomatic index to temperature. These differences were not present in a different population comparison between FB and BB cod. All populations had higher growth rates and food conversion efficiencies at warmer temperatures, and exhibited compensatory growth when temperature was increased. The results suggest relatively larger genetic differences between GB and GOM cod than between FB and BB cod, and indicate that the faster growth of southern populations in the wild is not due to a higher genetic capacity for growth rate.     

Minisatellite DNA fingerprints of Atlantic cod (Gadus morhua)

The human minisatellite probes 33.6 and 33.15 cross–hybridized to Hae III and Hinf I digested cod DNA, revealing complex fragment patterns in both Arctic and coastal cod. The DNA fingerprints were highly polymorphic, individual specific and stable in the germline. The potential applications of multi locus probes in cod research are discussed.     

Correlations between hemoglobin type and temperature preference of juvenile Atlantic cod Gadus morhua

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.     

Effects of temperature, hypoxia and activity on the metabolism of juvenile Atlantic cod

Standard metabolic rate (SMR), active metabolic rate (AMR) and critical oxygen saturation ( S_crit ) were measured in Atlantic cod Gadus morhua at 5, 10 and 15° C. The SMR was 35.5, 57.0 and 78.2 mg O₂ kg⁻¹ h⁻¹ and S_crit was 16.5, 23.2 and 30.3%, at 5, 10 and 15° C, respectively. Previously reported SMR for Atlantic cod from arctic waters at 4° C was twice that measured at 5° C in the present study. A possible intraspecific latitudinal difference in the SMR is discussed. The AMR was 146.6, 197.9 and 200.4 mg O₂ kg⁻¹ h⁻¹ and the critical swimming speed ( U_crit ) was 1 6, 1.7 and 1.9 at 5, 10 and 15° C, respectively. The maximum oxygen consumption was found to be associated with exercise, rather than recovery from exercise as previously reported in another Study of Cod metabolism.     

The effects of temperature and size on growth and mortality of cod larvae

The optimal temperatures for growth of four groups of hatchery-reared cod larvae (geometric mean weight: 73, 191, 249 and 251 μg), reared on rotifers at four or five constant temperatures between 4 and 16° C for 14, 12, 9 and 16 days were 9.7, 12.3, 12.7 and 13.4° C, respectively. The maximum growth rate also increased with size and was 6.5, 9.6, 11.7 and 11.3% day⁻¹ for the respective size groups. The optimal temperature for survival was 8.5–8.8° C for all size groups. The results indicate an opposite relationship between (1) size and optimal temperature for growth and (2) size and maximum growth rate of cod larvae, to that observed for juvenile and immature cod.     

Size-dependent mortality of juvenile Atlantic cod, estimated from recaptures of released reared cod and tagged wild cod

The recapture rate and survival of reared cod Gadus morhua L. (15–45 cm) released in fjords and open coastal areas of Norway increased with increasing size at release, while release area and season also had a significant influence on the results. High mortality soon after release reduced the number of released fish by more than two–thirds within 6 months, and reduced the potential commercial yield to a low level. In most cases the recapture yield was lower than the released biomass and the net yield increased little or not at all with increasing size at release. In western Norway high fishing mortality of young fish reduced the numbers of surviving fish rapidly and the growth overfishing reduced the potential yield still further. Natural mortality decreased with increasing size at release, and became low when the fish approached their size refuge, i.e. when they could not be eaten by their usual predators.