Die Reproduktion von Dorschen (Gadus morhua L. und Gadus morhua callarias L. ) in der Ostsee unter besonderer Berücksichtigung der Arkonasee. Teil I: Allgemeiner Verlauf des jährlichen Reifeprozesses und der Laichaktivitäten in den verschiedenen Gebieten

From 1993 to 2003 special surveys were conducted to analyse temporal and spatial aspects of the spawning activities of cod in the Baltic Sea. The ICES advice for cod in the Baltic Sea is based on yearly stock assessments of two separate stocks the western (SD 22 – 24) and the eastern stock (SD 25 – 32). Both stocks are characterized by different spawning seasons. The general progression of the maturity suggests that two types of development can be distinguished. In the western Baltic Sea (SD 22) the maturity development is temporal stable, with a relative short, main spawning period from March to April - spring spawner. In the Arkona Sea and Bornholm Basin the main spawning season starts in June and probably finishes in September - summer spawner. The proportion of spawners in summer is significantly higher than in spring in the Arkona Sea and Bornholm Basin. This again underlines the importance of the Arkona Sea for the reproduction success of the eastern Baltic cod stock. The analysis furthermore show, that the proportion of male spawners was significantly higher than the proportion of female spawners in all areas and the investigated months. This observation suggests that the reproductive success is only limited by the number of female cod which participates in the spawning process.

Die Reproduktion von Dorschen (Gadus morhua L. und Gadus morhua callarias L.) in der Ostsee unter besonderer Berücksichtigung der Arkonasee. Teil II: Statistische Analysen zum Anteil reproduktiv aktiver Dorsche in Bezug auf gebietsspezifische Unterschiede und Gemeinsamkeiten, sowie deren mögliche Ursachen.

Surveys were conducted to analyse temporal and spatial aspects of the spawning activities of cod in the Baltic Sea from 1993 to 2003. Part I of the article describes the general maturity development and the spawning activities in the western Baltic Sea, the Arkona Sea and the Bornholm Sea. The studies suggest that two types of spawner can be distinguished, the spring spawner in the western Baltic Sea and the summer spawner in the Arkona Sea and the Bornholm Basin. The descriptive analysis is supported by statistical studies presented in this article. It is shown, that the proportion of spawning individuals is significantly higher in summer than in spring in the ArkonaSea and the Bornholm Sea. Furthermore, it is shown that reproductive mixing of both types of spawner is possible in the Arkona Sea in May/June due to the hydrographical conditions in the Arkona Sea which were suitable in 8 of 10 years for successful reproduction of both Baltic cod stocks, and that spatial expansion of spawning activities of the eastern Baltic cod stock into the Arkona Sea is positive correlated with the size of spawning stock in the Bornholm Sea.

A simple approach for the estimation of food consumption from growth rates at different environmental conditions and its application to juvenile cod (Gadus morhua L.) of a fjordic sea loch on the west coast of Scotland

Abstract Growth and condition of fish are functions of available food and environmental conditions. This led to the idea of using fish as a “consumption sensor” for the measurement of food intake over a defined period of time. A bio-physical model for the estimation of food consumption was developed based on the von Bertalanffy model. Whereas some of the input variables of the model, the initial and final lengths and masses of a fish and the temperature within the time period considered can easily be measured, internal characteristics of the species have to be determined indirectly. Three internal parameters are used in the model: the maintenance consumption at 0°C, the temperature dependence of this consumption and the food efficiency, the percentage of the ingested food utilized. Estimates of the parameters for a given species can be determined by feeding experiments. Here, data from published feeding experiments on juvenile cod, Gadus morhua L., were used to validate the model. The average of the relative error for the food intake predicted by the model for individual fish was about 24 %, indicating that fish used the food with different efficiencies. However, grouping the fish according to size classes and temperature lowered the relative error of the predicted food intake for the group to typically 5 %. For a group containing all fish of the feeding experiment the relative prediction error was about 2 %. Zusammenfassung Wachstum und Kondition der Fische sind von der verfügbaren Nahrung und von Umweltbedingungen abhängig. Dies führte zur Idee, Fisch als „Konsum-Sensor“ für die Messung der Nahrungsaufnahme über einen definierten Zeitraum zu verwenden. Auf Grundlage des von Bertalanffy-Modells wurde ein bio-physikalisches Modell zur Schätzung der Futteraufnahme entwickelt. Während einige der Eingangsgrößen des Modells leicht gemessen werden können (Anfangs- und Endl��nge und -körpermasse der Fische und die Temperatur innerhalb des betrachteten Zeitraum), können interne Parameter der betrachteten Art nur indirekt bestimmt werden. Drei interne Parameter werden in dem Modell verwendet: Die Erhaltungskonsumtion bei 0° C, die Temperaturabhängigkeit dieser Rate und der Wirkungsgrad der Nahrung (der Anteil der Nahrung ,der aufgenommen und verwendet und nicht ungenutzt wieder ausgeschieden wird). Die Modellparameter für eine bestimmte Art können durch Fütterungsversuche bestimmt werden. Um das Modell zu validieren wurden Daten aus veröffentlichten Fütterungsversuchen mit juvenilen Kabeljau (Gadus morhua L.) verwendet. Modell und Wirklichkeit weichen in der Regel voneinander ab. Der durchschnittliche relative Fehler der durch das Modell vorhergesagten Nahrungsaufnahme betrug für Einzelfische etwa 24%, was darauf hinweist, dass einzelne Fisch die Nahrung mit unterschiedlichen Wirkungsgraden verwerten. Allerdings senkte die Gruppierung der Fische nach Größenklassen und Temperatur den relativen Vorhersagefehler für die Nahrungsaufnahme der Gruppe auf etwa 5%. Für alle Fische im Fütterungsversuch ist der relative Vorhersagefehler etwa 2%.

The effect of stress on egg quality in farmed Atlantic Cod (Gadus morhua L.)

Dissertação de mestrado, Aquacultura, Faculdade de Ciências e Tecnologia, Universidade do Algarve, 2011

The stress and metabolic responses of juvenile Atlantic cod Gadus morhua L. to an acute thermal challenge

Survival, oxygen consumption (M_O2), total plasma cortisol and glucose levels and gill heat-shock protein 70 (hsp70) expression were measured in 10 and 50 g juvenile Atlantic cod Gadus morhua during an acute temperature increase (2° C h⁻¹) to their critical thermal maximum. Ninety three per cent of the fish in both size classes survived to 24° C; however, mortality was 100% within 15 min of reaching this temperature. The M_O2 for both size classes increased significantly with temperature, reaching peak values at 22° C that were c. 2·8-fold those of control (10° C) fish. Resting plasma cortisol and glucose levels were lower in 10 g as compared to 50 g fish. Plasma glucose levels were highly variable in both size classes, and significant increases were only seen at >22° C for the 10 g fish. In contrast, plasma cortisol showed an exponential increase with temperature starting at 16° C in both size classes, and reached maximum levels at 22° C that were 19-fold (10 g fish) and 35-fold (50 g fish) higher than their respective control groups. Both the constitutive (73 kDa) and inducible (72 kDa) isoforms of hsp70 were detected in both size classes using the widely utilized mouse monoclonal antibody. Expression of these isoforms, however, did not change when Atlantic cod were exposed to elevated temperature, and the 72 kDa isoform was not detected using salmonid-specific antibodies. These results indicate that juvenile Atlantic cod are very sensitive to acute increases in water temperature. In addition, they (1) show that M_O2and plasma cortisol, but not plasma glucose or gill hsp 70 levels, are sensitive indicators of thermal stress in Atlantic cod and (2) support previous reports that the upper critical temperature for this species is 16° C.

Thermal acclimation to 4 or 10 degrees C imparts minimal benefit on swimming performance in Atlantic cod (Gadus morhua L.).

Thermal acclimation is frequently cited as a means by which ectothermic animals improve their Darwinian fitness, i.e. the beneficial acclimation hypothesis. As the critical swimming speed (U (crit)) test is often used as a proxy measure of fitness, we acclimated Atlantic cod (Gadus morhua) to 4 and 10 degrees C and then assessed their U (crit) swimming performance at their respective acclimation temperatures and during acute temperature reversal. Because phenotypic differences exist between different populations of cod, we undertook these experiments in two different populations, North Sea cod and North East Arctic cod. Acclimation to 4 or 10 degrees C had a minimal effect on swimming performance or U (crit), however test temperature did, with all groups having a 10-17% higher U (crit) at 10 degrees C. The swimming efficiency was significantly lower in all groups at 4 degrees C arguably due to the compression of the muscle fibre recruitment order. This also led to a reduction in the duration of "kick and glide" swimming at 4 degrees C. No significant differences were seen between the two populations in any of the measured parameters, due possibly to the extended acclimation period. Our data indicate that acclimation imparts little benefit on U (crit) swimming test in Atlantic cod. Further efforts need to identify the functional consequences of the long-term thermal acclimation process.

An examination of the metabolic processes underpinning critical swimming in Atlantic cod (Gadus morhua L.) using in vivo 31P-NMR spectroscopy.

Traditionally, critical swimming speed has been defined as the speed when a fish can no longer propel itself forward, and is exhausted. To gain a better understanding of the metabolic processes at work during a U(crit) swim test, and that lead to fatigue, we developed a method using in vivo (31)P-NMR spectroscopy in combination with a Brett-type swim tunnel. Our data showed that a metabolic transition point is reached when the fish change from using steady state aerobic metabolism to non-steady state anaerobic metabolism, as indicated by a significant increase in inorganic phosphate levels from 0.3+/-0.3 to 9.5+/-3.4 mol g(-1), and a drop in intracellular pH from 7.48+/-0.03 to 6.81+/-0.05 in muscle. This coincides with the point when the fish change gait from subcarangiform swimming to kick-and-glide bursts. As the number of kicks increased, so too did the Pi concentration, and the pH(i) dropped. Both changes were maximal at U(crit). A significant drop in Gibbs free energy change of ATP hydrolysis from -55.6+/-1.4 to -49.8+/-0.7 kJ mol(-1) is argued to have been involved in fatigue. This confirms earlier findings that the traditional definition of U(crit), unlike other critical points that are typically marked by a transition from aerobic to anaerobic metabolism, is the point of complete exhaustion of both aerobic and anaerobic resources.

The swimming kinematics of larval Atlantic cod, Gadus morhua L., are resilient to elevated seawater pCO2

Kinematics of swimming behavior of larval Atlantic cod, aged 12 and 27 days post-hatch (dph) and cultured under three pCO2 conditions (control-370, medium-1800, and high-4200 µatm) from March to May 2010, were extracted from swim path recordings obtained using silhouette video photography. The swim paths were analyzed for swim duration, distance and speed, stop duration, and horizontal and vertical turn angles to determine whether elevated seawater pCO2-at beyond near-future ocean acidification levels-affects the swimming kinematics of Atlantic cod larvae. There were no significant differences in most of the variables tested: the swimming kinematics of Atlantic cod larvae at 12 and 27 dph were highly resilient to extremely elevated pCO2 levels. Nonetheless, cod larvae cultured at the highest pCO2 concentration displayed vertical turn angles that were more restricted (median turn angle, 15°) than larvae in the control (19°) and medium (19°) treatments at 12 dph (but not at 27 dph). Significant reduction in the stop duration of cod larvae from the high treatment (median stop duration, 0.28 s) was also observed compared to the larvae from the control group (0.32 s) at 27 dph (but not at 12 dph). The functional and ecological significance of these subtle differences are unclear and, therefore, require further investigation in order to determine whether they are ecologically relevant or spurious.

Effects of ocean acidification on the calcification of otoliths of larval Atlantic cod Gadus morhua

The growth and development of the aragonitic CaCO3 otoliths of teleost fish could be vulnerable to processes resulting from ocean acidification. The potential effects of an increase in atmospheric CO2 on the calcification of the otoliths were investigated by rearing Atlantic cod Gadus morhua L. larvae in 3 pCO2 concentrations-control (370 µatm), medium (1800 µatm) and high (4200 µatm)-from March to May 2010. Increased otolith growth was observed in 7 to 46 d post hatch (dph) cod larvae at elevated pCO2 concentrations. The sagittae and lapilli were usually largest in the high pCO2 treatment followed by the medium and control treatments. The greatest difference in mean otolith surface area (normalized to fish length) was for sagittae at 11 dph, with medium and high treatments being 46 and 43% larger than the control group, respectively. There was no significant pCO2 effect on the shape of the otoliths nor were there any trends in the fluctuating asymmetry, defined as the difference between the right and left sides, in relation to the increase in otolith growth from elevated pCO2.

Hemoglobin genotype has minimal influence on the physiological response of juvenile atlantic cod (Gadus morhua) to environmental challenges

Hemoglobin (Hb) polymorphism in cod is associated with temperature‐related differences in biogeographical distribution, and several authors have suggested that functional characteristics of the various hemoglobin isoforms (HbIs) directly influence phenotypic traits such as growth rate. However, no study has directly examined whether Hb genotype translates into physiological differences at the whole animal level. Thus, we generated a family of juvenile Atlantic cod consisting of all three main Hb genotypes (HbI‐1/1, HbI‐2/2, and HbI‐1/2) by crossing a single pair of heterozygous parents, and we compared their metabolic and cortisol responses to an acute thermal challenge (10°C to their critical thermal maximum [CTM] or 22°C, respectively) and tolerance of graded hypoxia. There were no differences in routine metabolism (at 10°C), maximum metabolic rate, metabolic scope, CTM (overall mean 22.9° ± 0.2°C), or resting and poststress plasma cortisol levels among Hb genotypes. Further, although the HbI‐1/1 fish grew more (by 15%–30% during the first 9 mo) when reared at 10° ± 1°C and had a slightly enhanced hypoxia tolerance at 10°C (e.g., the critical O2 levels for HbI‐1/1, HbI‐2/2, and HbI‐1/2 cod were 35.56% ± 1.24%, and 40.20% ± 1.99% air saturation, respectively), these results are contradictory to expectations based on HbI functional properties. Thus, our findings (1) do not support previous assumptions that growth rate differences among cod Hb genotypes result from a more efficient use of the oxygen supply—that is, reduced standard metabolic rates and/or increased metabolic capacity—and (2) suggest that in juvenile cod, there is no selective advantage to having a particular Hb genotype with regards to the capacity to withstand ecologically relevant environmental challenges.

In situ cardiac performance of Atlantic cod (Gadus morhua) at cold temperatures: long-term acclimation, acute thermal challenge and the role of adrenaline

The resting and maximum in situ cardiac performance of Newfoundland Atlantic cod (Gadus morhua) acclimated to 10, 4 and 0°C were measured at their respective acclimation temperatures, and when acutely exposed to temperature changes: i.e. hearts from 10°C fish cooled to 4°C, and hearts from 4°C fish measured at 10 and 0°C. Intrinsic heart rate (f(H)) decreased from 41 beats min(-1) at 10°C to 33 beats min(-1) at 4°C and 25 beats min(-1) at 0°C. However, this degree of thermal dependency was not reflected in maximal cardiac output (Q(max) values were ~44, ~37 and ~34 ml min(-1) kg(-1) at 10, 4 and 0°C, respectively). Further, cardiac scope showed a slight positive compensation between 4 and 0°C (Q(10)=1.7), and full, if not a slight over compensation between 10 and 4°C (Q(10)=0.9). The maximal performance of hearts exposed to an acute decrease in temperature (i.e. from 10 to 4°C and 4 to 0°C) was comparable to that measured for hearts from 4°C- and 0°C-acclimated fish, respectively. In contrast, 4°C-acclimated hearts significantly out-performed 10°C-acclimated hearts when tested at a common temperature of 10°C (in terms of both Q(max) and power output). Only minimal differences in cardiac function were seen between hearts stimulated with basal (5 nmol l(-1)) versus maximal (200 nmol l(-1)) levels of adrenaline, the effects of which were not temperature dependent. These results: (1) show that maximum performance of the isolated cod heart is not compromised by exposure to cold temperatures; and (2) support data from other studies, which show that, in contrast to salmonids, cod cardiac performance/myocardial contractility is not dependent upon humoral adrenergic stimulation.

Use of Otolith Strontium:Calcium Ratios for Hindcasting Larval Cod Gadus Morhua Distributions Relative to Water Masses on Georges Bank

The concentration ratios of strontium to calcium in laboratory-reared larval cod otoliths are shown to be related to the water temperature (T) at the time of otolith precipitation. This relationship is curvilinear, and is best described by a simple exponential equation of the form (Sr/Ca x 1000 = a exp(-T/b). We show that when Sr/Ca elemental analyses are related to the daily growth increments in the larval otoliths, relative temperature histories of individual field-caught larvae can be reconstructed from the egg stage to the time of capture. We present preliminary examples of how such reconstructed temperature histories of Atlantic cod Gadus morhua larvae, collected on Georges Bank during April and May 1993, may be interpreted in relation to the broad-scale larval distributions and the hydrography of the Bank.