Reconstructed 3D models of digestive organs of developing Atlantic cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) larvae

Six 3D models of the digestive system during ontogeny were reconstructed from histological sections of Atlantic cod larvae. The 3D models clearly visualize the following features: folding of the gut rotation; subdivision of digestive tract into foregut, midgut, and hindgut by sphincters; development of stomach and pyloric caeca from 39 dph; location of entrances of bile and pancreatic ducts in the medial plane of the anterior midgut; ontogeny of pancreas from a compact organ to an elongated and branched (but not diffuse) organ along the posterior midgut; one dominant islet of Langerhans until 39 dph and several smaller satellite islets also visible from 53 dph; the relatively large volume of the anterior midgut that probably increases residence time of ingested food mixing with secretions from pancreas and bile. Calculated volumes of each digestive organ demonstrate allometric changes during ontogeny. Interactive 3D models are available as QuickTime format downloadable files.     

Relation of growth and condition with the <Emphasis Type="Italic">Pan</Emphasis> I locus in Atlantic cod (<Emphasis Type="Italic">Gadus morhua</Emphasis> L.) around Iceland

Growth and condition (both somatic and hepatosomatic index) of Atlantic cod spawning at different locations around Iceland was studied in relation to the Pan I locus. South of Iceland cod carrying the Pan I^AB and Pan I^BB genotypes were more frequent while cod carrying the Pan I^AA genotype was more frequent north of Iceland. Differences in growth were detected between cod spawning at different areas around Iceland. Cod spawning south of Iceland grew faster than cod spawning north of Iceland. Differences in growth rate were also observed among cod carrying different Pan I genotypes within a spawning area. The least frequent Pan I genotype expressed the highest growth in both south and north of Iceland. Cod carrying the Pan I^AA grew fastest at spawning locations south of Iceland, while cod carrying the Pan I^BB genotype grew fastest in north of Iceland. A consistent relationship between condition and the different Pan I genotypes was also observed in all the areas. Cod carrying the Pan I^AA expressed the highest somatic condition and the lowest hepatosomatic index. Together, these results indicate that the relation of growth and condition with the Pan I locus is more complicated than earlier thought and is likely to be influenced by other factors, like size-selective fishing and food supply.     

Foraging behaviour of larval cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) at low light intensities

The ability to forage at low light intensities can be of great importance for the survival of fish larvae in a pelagic environment. Three-dimensional silhouette imaging was used to observe larval cod foraging and swimming behaviour at three light intensities (dusk ~1.36 × 10⁻³ W/m², night ~1.38 × 10⁻⁴ W/m² and darkness ~3.67 × 10⁻⁶ W/m²) at 4 different ages from 6 to 53 days post-hatch (dph). At 6 dph, active pursuit of prey was only observed under dusk conditions. Attacks, and frequent orientations, were observed from 26 dph under night conditions. This was consistent with swimming behaviour which suggested that turn angles were the same under dusk and night conditions, but lower in darkness. Cod at 53 dph attacked prey in darkness and turn angles were not different from those under other light conditions. This suggests that larvae are still able to feed at light intensities of 3.67 × 10⁻⁶ W/m². We conclude that larval cod can maintain foraging behaviour under light intensities that correspond to night-time at depths at which they are observed in the field, at least if they encounter high-density patches of prey such as those that they would encounter at thin layers or fronts.     

Fluctuating asymmetry and nutritional condition of Baltic cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) larvae

The level of fluctuating asymmetry (FA), defined as random deviations from perfect bilateral symmetry, is assumed to reflect the developmental instability (DI) of an organism. Because environmental and genetic stress may increase DI, FA has been used to assess the level of stress experienced by, for example, fish. In this study, left–right asymmetry of lapillar otoliths was related to nutritional condition as estimated from RNA/DNA ratios, in order to investigate the utility of FA to detect feeding-related stress in Baltic cod, Gadus morhua L., larvae. Cod larvae in intermediate and good nutritional condition showed similar values of FA, and these were more symmetric than for larvae in poor condition. As increased levels of FA were restricted to larvae in a condition comparable to that of larvae experiencing at least 3 days of starvation in laboratory experiments, it is suggested that FA is an insensitive indicator of short-term feeding success of larval cod. However, FA can be used to reveal severely starved larva populations and probably also populations that have been subjected to prolonged sub-optimal feeding conditions.     

Modelling temporal and spatial variability in tag reporting-rates for Newfoundland cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>)

Mark–recapture experiments can be used to estimate the exploitation rate of a fishery; however, the estimate is influenced by the tag reporting-rate by the fishers. We present two methods to estimate the reporting rates in high/low reward ($100 and $10 CAD respectively) long-term cod tagging experiments. We fit two binomial logistic mixed-effect models, one with temporal auto-correlation in the reporting-rate year-effects and one with independent year-effects. We estimate reporting-rates separately for recreational and commercial fishers, and test for spatial variation using fixed-effects for spatial regions. Due to the complexity of the fishery, our models account for factors such as recapture-fishery type, fish-size and time-at-liberty. Our results indicate that the recreational fishers reporting-rate was constant at 0.51 across all regions and years. The commercial fishery showed more spatial and temporal variation, with the reporting-rates estimates lying between 0.67 and 0.87 for the independent year-effect model, and between 0.57 and 0.84 for the random walk model. Furthermore, we assessed the model performance as well as the coverage probability of nominal 95 % confidence intervals using simulations. We found that the models performed adequately; however, the nominal 95 % confidence intervals tended to be too narrow.     

Diet of 0-group stages of capelin (<Emphasis Type="Italic">Mallotus villosus</Emphasis>), herring (<Emphasis Type="Italic">Clupea harengus</Emphasis>) and cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) during spring and summer in the Barents Sea

Recruitment of capelin in the Barents Sea fail when juvenile herring and cod are abundant and the potential for feeding competition of wild sympatric capelin and herring larvae and small cod juveniles were investigated. The frequency of gut evacuation after capture of capelin larvae were also studied in mesocosms. Small capelin larvae (<35 mm length) fed on small prey including phytoplankton, invertebrate eggs and nauplii, bivalves, other invertebrate larvae and small copepods. Calanus copepodites were only observed in large capelin larvae (>26 mm length). Calanus copepodites were the major food sources for contemporary herring larvae (25–35 mm length) and Calanus and euphausiids were the major prey for small juvenile herring (37–60 mm length) and cod (18–40 mm length). Capelin larvae reared in mesocosms evacuated the guts shortly after capture. Capelin larvae had a smaller mouth and fed on smaller prey than herring and cod of the same length. This implies that the small capelin larvae, in contrast to sympatric small herring and cod, are not tightly linked to the food chain involving Calanus and euphausiids. Thus, exploitative competition between capelin larvae and planktivorous fish that rely on Calanus and euphausiids in the Barents Sea may be relaxed.     

Temperature and developmental plasticity during embryogenesis in the Atlantic cod<Emphasis Type="Italic"> Gadus morhua</Emphasis> L.

Atlantic cod (Gadus morhua L.) embryos were reared at 4 °C, 7 °C, and 10 °C, and the relative timing of developmental events was characterized, with particular reference to myotomal muscle. Embryos started to feed at an apparently equivalent stage of development, so comparisons were made between temperature groups on the basis of percentage of time to first feeding and somite stage. No differences were found in the time of hatching or timing of appearance of the otic placode, unpaired median fin fold, gut lumen, otic vesicle, lens of the eye, otoliths, first muscular contractions, swim bladder, and hindgut, or in the rate of development of somites, myotubes, myofibrils, and acetylcholinesterase activity over the temperature range studied. In contrast, closure of the blastopore occurred late with respect to segmentation at higher temperatures, at the 3-somite, 10-somite, and 12-somite stages at 4 °C, 7 °C, and 10 °C respectively. Muscle cellularity was also markedly altered in the 10 °C group relative to the 4 °C and 7 °C groups. Larvae reared at 10 °C had significantly more (+14%) deep white fibers at hatch (P <0.001), whereas numbers of superficial red fibers remained unchanged. It is suggested that differences in muscle cellularity might be related to changes in the relative timing of epiboly, through differential proliferation of presomitic myogenic cells and/or their relative exposure to inductive signals.     

Isolation of Atlantic cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) nursery areas

This study examines the degree of isolation among 0-group Atlantic cod in a region where a number of resident adult congregations have been found. The location of key nursery areas in Scottish coastal waters was identified from trawl surveys of settled 0-group cod around the Scottish coast. Segregation among four of these key areas; within the Clyde, Shetland Isles, Moray Firth and Buchan was investigated using otolith elemental composition. Isotope concentrations in the otoliths of cod were analysed by solution-based inductively coupled plasma mass spectrometry of the whole otolith. Discriminant analyses showed a clear separation between nursery areas based on the concentrations of three elements (manganese, barium and magnesium). The concentration of these elements was not affected by the age or size of individuals. Classification accuracy exceeded 90% for all four sites. Evidence of temporal persistence in the relative spatial differences in elemental concentrations between two sites for 2 year-classes was found. The results suggest limited exchange between sites during some period of the early life history. The significance of this limited exchange is discussed in relation to recent evidence for adult fidelity and the utility of closed areas for protecting nursery grounds.     

The response of Atlantic cod, <Emphasis Type="Italic">Gadus morhua</Emphasis>, to progressive hypoxia: fish swimming speed and physiological stress

Atlantic cod, Gadus morhua, were exposed to a progressive stepwise decline in water oxygen pressure Fish swimming speed and indicators of primary and secondary stress (e.g. blood cortisol and lactate) were measured to assess whether a severe shift in physiological homeostasis (i.e. stress) preceded any change in behaviour or vice versa. Swimming speed increased by 18% when was reduced rapidly from 19.9 kPa to 13.2 kPa and was interpreted as an initial avoidance response. However, swimming speed was reduced by 21% at a moderate level of steady (8.4 kPa) and continued to drop by 41% under progressively deep hypoxia (4.3 kPa). Elevations in plasma cortisol and blood lactate indicated major physiological stress but only at 4.3 kPa, which corresponds to the critical oxygen tension of this species. We propose that the drop in speed during hypoxia aids to offset major stress and is adaptive for the survival of cod in extensive areas of low oxygen.     

The hypoxia avoidance behaviour of juvenile Atlantic cod (<Emphasis Type="Italic">Gadus morhua</Emphasis> L.) depends on the provision and pressure level of an O<Subscript>2</Subscript> refuge

The frequency of low O₂ (hypoxia) has increased in coastal marine areas but how fish avoid deleterious water masses is not yet clear. To assess whether the presence and oxygen pressure (PO₂) level of an O₂ refuge affects the hypoxia avoidance behaviour of fish, individual Atlantic cod (Gadus morhua L.) were exposed to a range of O₂ choices in a 2-way choice chamber at 11.4°C over two different experiments. Cod in the first experiment were allowed access to a fixed O₂ refuge (fully air-saturated seawater) whilst oxygen pressure (PO₂) on the other side was reduced in steps to a critically low level, i.e. 4.3 kPa—a point where cod can no longer regulate O₂ consumption. Under these conditions, cod did not avoid any level of hypoxia and fish swimming speed also remained unchanged. In contrast, strong avoidance reactions were exhibited in a second experiment when fish were again exposed to 4.3 kPa but the safety, i.e. PO₂, of the refuge was reduced. Fish not only spent less time at 4.3 kPa as a result of fewer sampling visits but they also swam at considerably slower speeds. The presence of an avoidance response was thus strongly related to refuge PO₂ and it is unlikely that cod, and possibly other fish species, would enter low O₂ to feed in the wild if a sufficiently safe O₂ refuge was not available. It is therefore hypothesized that the feeding range of fish may be heavily compressed if hypoxia expands and intensifies in future years.     

Precision and accuracy of stable isotope signals in otoliths of pen-reared cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) when sampled with a high-resolution micromill

A high-resolution sampling technique for stable-isotope measurements in otoliths is now available, but little is known about the precision and accuracy of the results when using this technique. Otolith samples of pen-reared cod (Gadus morhua L.) with well-defined temperature history were obtained using this high-resolution sampling technique. Samples from different-aged cod otoliths were collected in two yearly increments corresponding to the calendar years 1994 and 1995, and the stable-oxygen (¹⁸O) and -carbon (¹³C) isotope compositions of the samples were measured. Otolith ¹⁸O values had a clear seasonal pattern in response to the seasonal water temperature. Otolith material deposited in the same calendar years of different-aged fish showed similar ¹⁸O values. High precision of the temperature estimate using otolith ¹⁸O values was obtained when comparing results from different otoliths. The accuracy of the temperature estimates, relative to measured mean monthly water temperature, was also high, but reduced sampling resolution in the otoliths significantly reduced the accuracy due to attenuation of the ¹⁸O signals. We found that the otolith ¹³C values had a cyclic pattern, roughly in phase with the ¹⁸O values. This was most likely caused by temperature-increased metabolism. Indications of age-specific otolith ¹³C values are also presented.Communicated by M. Kühl, Helsingør     

Genetic analysis of tuna populations,<Emphasis Type="Italic"> Thunnus thynnus thynnus</Emphasis> and<Emphasis Type="Italic"> T</Emphasis>.<Emphasis Type="Italic"> alalunga</Emphasis>

The genetic population structures of Atlantic northern bluefin tuna ( Thunnus thynnus thynnus) and albacore ( T. alalunga) were examined using allozyme analysis. A total of 822 Atlantic northern bluefin tuna from 18 different samples (16 Mediterranean, 1 East Atlantic, 1 West Atlantic) and 188 albacore from 5 samples (4 Mediterranean, 1 East Atlantic) were surveyed for genetic variation in 37 loci. Polymorphism and heterozygosity reveal a moderate level of genetic variability, with only two highly polymorphic loci in both Atlantic northern bluefin tuna ( FH* and SOD- 1*) and albacore ( GPI- 3* and XDH*). The level of population differentiation found for Atlantic northern bluefin tuna and albacore fits the pattern that has generally been observed in tunas, with genetic differences on a broad rather than a more local scale. For Atlantic northern bluefin tuna, no spatial or temporal genetic heterogeneity was observed within the Mediterranean Sea or between the East Atlantic and Mediterranean, indicating the existence of a single genetic grouping on the eastern side of the Atlantic Ocean. Very limited genetic differentiation was found between West Atlantic and East Atlantic/Mediterranean northern bluefin tuna, mainly due to an inversion of SOD- 1* allele frequencies. Regarding albacore, no genetic heterogeneity was observed within the Mediterranean Sea or between Mediterranean and Azores samples, suggesting the existence of a single gene pool in this area.     

Histological investigation of the reproductive cycles of the limpets <Emphasis Type="Italic">Patella </Emphasis><Emphasis Type="Italic">vulgata</Emphasis> and <Emphasis Type="Italic">Patella ulyssiponensis</Emphasis>

This study devised a staging system for, and monitored, the gonad development of the limpet species Patella vulgata and Patella ulyssiponensis on the South West coast of Ireland using histological techniques. Maturation began in the males of both species in January and in the females it began in March. There was no statistical difference in gonad development between sexes and between species. Spawning in the male P. vulgata occurred from September to December 2003 and in September and October 2004. In female P. vulgata spawning occurred from October to December 2003, no spawning of females was observed in 2004. In male P. ulyssiponensis spawning occurred in November and December 2003 and from September 2004 to December 2004. Spawning was observed from November 2003 to January 2004 and in September 2004 in female P. ulyssiponensis. Sex ratios also varied between the species and between months sampled. Nevertheless more males were observed in both species.     

Heart rate reflects osmostic stress levels in two introduced colonial ascidians <Emphasis Type="Italic">Botryllus schlosseri</Emphasis> and <Emphasis Type="Italic">Botrylloides</Emphasis><Emphasis Type="Italic">violaceus</Emphasis>

The influence of abiotic factors on the establishment and success of invasive species is often difficult to determine for most marine ecosystems. However, examining this relationship is critical for predicting the spread of invasive species and predicting which habitats will be most vulnerable to invasion. Here we examine the mortality and physiological sensitivity to salinity of adult colonies of the colonial ascidians Botryllus schlosseri and Botrylloides violaceus. Adult colonies of each species were exposed to abrupt changes in salinity (5, 10, 15, 20, 25, 30 psu) in the laboratory. Salinity ranges used in the laboratory corresponded with those of the field distributions of B. violaceus and B. schlosseri in the Great Bay Estuary, NH. Heart rate was used as a proxy for health to assess the condition of individual colonies. Heart rates were monitored daily for approximately 2 weeks. Results revealed that both species experienced 100% mortality after 1 day at 5 psu and that their heart rates declined with decreasing salinity. Heart rates of B. schlosseri remained consistent between 15 and 30 psu and slowed at 10 psu. Heart rates of B. violaceus remained constant between 20 and 30 psu, but slowed at 15 psu. These laboratory results corresponded to the distribution of these species in estuaries, indicating salinity is a key factor in the distribution and dominance of B. schlosseri and B. violaceus among coastal and estuarine sites. Furthermore, physiological differences to salinity were found between colonies of B. schlosseri in the Venetian Lagoon and colonies in Portsmouth Harbor, suggesting adaptation to environmental variables.     

Auditory defence in the peacock butterfly (<Emphasis Type="Italic">Inachis io</Emphasis>) against mice (<Emphasis Type="Italic">Apodemus flavicollis</Emphasis> and <Emphasis Type="Italic">A. sylvaticus</Emphasis>)

Morphological and behavioural traits can serve as anti-predator defence either by reducing detection or recognition risks, or by thwarting initiated attacks. The latter defence is secondary and often involves a ‘startle display’ comprising a sudden release of signals targeting more than one sensory modality. A suggested candidate for employing a multimodal defence is the peacock butterfly, Inachis io, which, by wing-flicking suddenly, produces sonic and ultrasonic sounds and displays four large eyespots when attacked. The eyespots make small birds retreat, but whether the sounds produced thwart predator attacks is largely unknown. Peacocks hibernate as adults in dark wintering sites and employ their secondary defence upon encounter with small rodent predators during this period. In this study, we staged predator–prey encounters in complete darkness in the laboratory between wild mice, Apodemus flavicollis and Apodemus sylvaticus, and peacocks which had their sound production intact or disabled. Results show that mice were more likely to flee from sound-producing butterflies than from butterflies which had their sound production disabled. Our study presents experimental evidence that the peacock butterfly truly employs a multimodal defence with different traits targeting different predator groups; the eyespots target birds and the sound production targets small rodent predators.     

Patterns of gamete incompatibility between the blue mussels<Emphasis Type="Italic"> Mytilus edulis</Emphasis> and<Emphasis Type="Italic"> M</Emphasis>.<Emphasis Type="Italic"> trossulus</Emphasis>

Previous research on gametic incompatibility in marine invertebrates suggests that for highly dispersive marine invertebrate species, barriers to fertilization among closely related taxa are often incomplete and sometimes asymmetric. The nature of these barriers can dramatically affect the patterns of gene flow and genetic differentiation between species, and thus speciation. Blue mussels, in the genus Mytilus, are genetically distinct in allopatry yet hybrids are present wherever any two species within the group co-occur. The present study sampled M. edulis (L.) and M. trossulus (Gould) in May and June 2001 from the East Bay section of Cobscook Bay, Maine, USA (latitude 44°56′30″N; longitude 67°07′50″W), where the two species are sympatric. Gamete incompatibility was investigated in a series of laboratory fertilizations carried out in July 2001. The proportion of fertilized eggs typically exceeded 80% at sperm concentrations of 10³–10⁴ ml^?1 among intraspecific matings (n=18), but was <30% even at sperm concentrations in excess of 10⁵–10⁶ ml^?1 for interspecific matings (n=13). Further analysis indicated that approximately 100- to 700-fold higher sperm concentrations were required to achieve 20% fertilization in interspecific matings relative to intraspecific matings, indicating strong barriers to interspecific fertilization. The proportion of fertilized eggs did not follow this general pattern in all matings, however. The eggs from two (out of five) M. edulis females were almost as receptive to M. trossulus sperm as they were to M. edulis sperm. In contrast, the eggs from all M. trossulus females (n=3) were unreceptive to M. edulis sperm, suggesting that fertilization barriers between these species may be asymmetric. Given the experimental design employed in this study, the results are also consistent with a strong maternal or egg effect on the level of interspecific gamete compatibility in M. edulis.     

Feeding selectivities of the marine cladocerans<Emphasis Type="Italic"> Penilia avirostris</Emphasis>,<Emphasis Type="Italic"> Podon intermedius</Emphasis> and<Emphasis Type="Italic"> Evadne nordmanni</Emphasis>

We conducted grazing experiments with the three marine cladoceran genera Penilia, Podon and Evadne, with Penilia avirostris feeding on plankton communities from Blanes Bay (NW Mediterranean, Spain), covering a wide range of food concentrations (0.02–8.8 mm³ l^–1, plankton assemblages grown in mesocosms at different nutrient levels), and with Podon intermedius and Evadne nordmanni feeding on the plankton community found in summer in Hopavågen Fjord (NE Atlantic, Norway, 0.4 mm³ l^–1). P. avirostris and P. intermedius showed bell-shaped grazing spectra. Both species reached highest grazing coefficients at similar food sizes, i.e. when the food organisms ranged between 15 and 70 µm and between 7.5 and 70 µm at their longest linear extensions, respectively. E. nordmanni preferred organisms of around 125 µm, but also showed high grazing coefficients for particles of around 10 µm, while grazing coefficients for intermediate food sizes were low. Lower size limits were >2.5 µm, for all cladocerans. P. avirostris showed upper food size limits of 100 µm length (longest linear extension) and of 37.5 µm particle width. Upper size limits for P. intermedius were 135 µm long and 60 µm wide; those for E. nordmanni were 210 µm long and 60 µm wide. Effective food concentration (EFC) followed a domed curve with increasing nutrient enrichment for P. avirostris; maximum values were at intermediate enrichment levels. The EFC was significantly higher for P. intermedius than for E. nordmanni. With increasing food concentrations, the clearance rates of P. avirostris showed a curvilinear response, with a narrow modal range; ingestion rates indicated a rectilinear functional response. Mean clearance rates of P. avirostris, P. intermedius and E. nordmanni were 25.5, 18.0 and 19.3 ml ind.^–1 day^–1, respectively. Ingestion rates at similar food concentrations (0.4 mm³ l^–1) were 0.6, 0.8 and 0.9 g C ind.^–1 day^–1.Communicated by O. Kinne, Oldendorf/Luhe