Susceptibility of Atlantic herring and plaice larvae to predation by juvenile cod and herring at two constant temperatures

Predation encounters were staged in the laboratory to compare prey responsiveness, predator error rate, and predator capture success for juvenile cod Gadus morhua (a suction feeder) and herring Clupea harengus (a biting predator) preying on herring and plaice Pleuronectes platessa larvae. Trials were conducted at near natural temperature extremes for the larvae (8 and 13°C) to assess the importance of water temperature to the interaction. Herring larvae were significantly more responsive to attacks by herring than were plaice larvae (5·7 vs 0'0%). The two prey species were equally responsive to attacks by cod (2–6 vs 10%). Cod caught 91% of herring larvae attacked and juvenile herring caught 87%. Cod were successful in 96% of attacks on plaice, but juvenile herring caught significantly fewer (83%) plaice larvae. For each predator species, capture success did not vary significantly with prey species. Overall capture success for herring was significantly lower than that for cod. Responsiveness of herring larvae to attacks by juvenile herring increased with temperature, but predator error rate and capture success were not altered by water temperature.     

New data on trematodes of the families Opecoelidae and Accacoeliidae from fishes in Atlantic ocean and its seas

The additions concerning the morphological characters and measurements of Anomalotrema koiae, Helicometra fasciata, Accacladocoelium macrocotyle, A. petasiporum, and Paraccacladium jamiesoni have been made. Podocotyle atomon (Rud., 1802) in Gaevskaya, Umnova, 1977 described from Sebastes marinus from the northwest Atlantic is considered as Anomalotrema koiae. Helicometra fasciata is found in Mugilidae for the first time. Accacladocoelium macrocotyle and A. petasiporum are found in the new region--the Azores.     

Cholecystokinin mRNA in Atlantic herring, Clupea harengus--molecular cloning, characterization, and distribution in the digestive tract during the early life stages

The mRNA of the peptide hormone cholecystokinin (CCK) was isolated from juvenile Atlantic herring, Clupea harengus, by RT-PCR. The open reading frame encodes a 137 amino acid-long precursor protein. The peptide sequence of herring CCK-8, DYMGWMDF, is identical to that of higher vertebrates and elasmobranchs, and contains methionine in the sixth position from the C-terminus, which has not been reported previously in teleosts. Expression analysis by in situ hybridization shows that positive endocrine-like cells were mainly located in the pyloric caeca and to a less extent in the rectum of the juvenile. A few positive cells were also found in the pyloric portion of the stomach and the intestine. CCK cells were present in all the larvae examined from the day of hatching onwards. Although the CCK cells were scattered throughout the whole midgut, no signals were detected in either the foregut or the hindgut. Since herring larvae have a straight gut, the distribution pattern of CCK cells seems to be reflected in the anatomy of the gut.     

Drinking and water absorption by the larvae of herring (Clupea harengus) and turbot (Scophthalmus maximus)

The efficiencies of water absorption from the guts of the larvae of herring ( Clupea harengus L.) and turbot ( Scophthalmus maximus L.) were estimated by two methods, The first method was based on the differences in the rates of accumulation, by drinking, and clearance from the gut of radiolabelled inert markers. The second method used the equilibrium level of radioactivity in the larvae to measure the concentration of the markers in the gut above background as water is absorbed from the gut. Water absorption efficiencies for herring larvae drinking sea water were estimated to be 77% using both methods. When external salinity was reduced to 50% sea water, drinking rates and water absorption efficiency in herring larvae fell substantially.
Estimates of water absorption efficiency of seawater-adapted turbot larvae were similar (71–84%) to that of herring using both methods. Although temperature had a marked effect on both the rate of drinking and water absorption, there was no significant thermal effect on the efficiency of absorption from the guts of turbot larvae. The limitations of the techniques and the implications of the estimates in terms of water balance in fish larvae are discussed.     

Is the marginal otolith increment width a reliable recent growth index for larval and juvenile herring?

Marginal otolith increment width analysis was performed on field‐collected larval and juvenile spring‐spawned herring Clupea harengus that experienced variable feeding conditions and high temperatures that were above the optimum for growth. Although drastic zooplankton biomass reduction had a significant effect on increment width, a delay of a few days in the otolith response was observed. More importantly, a very clear, positive temperature effect on marginal increment width was demonstrated in fish characterized by temperature independent somatic growth. These results indicate that under natural conditions it may be impossible to distinguish increment width changes related to variation in feeding conditions from changes caused by temperature fluctuations. Therefore, it was concluded that marginal otolith increment width analysis could not be used as a recent growth index ( I _G) for herring larvae and juveniles exposed to drastic temperature fluctuations. The implication of these results is significant not only for the use of marginal increments as a recent growth index, but also if growth rate backcalculation is to be used as a research method.     

Temperature-driven changes in early life-history stages influence the Gulf of Riga spring spawning herring (<Emphasis Type="Italic">Clupea harengus m</Emphasis>.) recruitment abundance

Processes occurring during early life-history stages influence the year-class abundance of marine fish. We found that the abundance of 1-year-old spring spawning herring is statistically significantly determined by the number of post-flexion herring larvae in the Gulf of Riga (Baltic Sea). The abundance of consecutive developmental stages of larvae: yolk-sac, pre-flexion, flexion and post-flexion strongly correlated with each other, indicating that factors which already influence the yolk-sac stage are important in determining the abundance of post-flexion herring larvae. Winter air temperature before spawning determined the timing of maximum abundance of pre-flexion herring larvae, but not their main prey: copepod nauplii, implying that different mechanisms governing major preconditions for the formation of year-class strength. The abundance of post-flexion larvae displayed a potential dome-shaped relationship with sea surface temperature experienced after hatching. We suggest that increased summer temperatures, which exceed the physiological optimum negatively, affect the survival of post-flexion herring larvae. Overall, future climate warming poses an additional risk to larval herring survival and this may lead to a reduction in those herring stock which rely on recruitment from shallow coastal areas.     

Larval and juvenile Pacific herring Clupea pallasii are not susceptible to infectious hematopoietic necrosis under laboratory conditions

Infectious hematopoietic necrosis (IHN) leads to periodic epidemics among certain wild and farmed fish species of the Northeast (NE) Pacific. The source of the IHN virus (IHNV) that initiates these outbreaks remains unknown; however, a leading hypothesis involves viral persistence in marine host species such as Pacific herring Clupea pallasii. Under laboratory conditions we exposed specific pathogen-free (SPF) larval and juvenile Pacific herring to 10(3) to 10(4) plaque-forming units (pfu) of IHNV ml(-1) by waterborne immersion. Cumulative mortalities among exposed groups were not significantly different from those of negative control groups. After waterborne exposure, IHNV was transiently recovered from the tissues of larvae but absent in tissues of juveniles. Additionally, no evidence of viral shedding was detected in the tank water containing exposed juveniles. After intraperitoneal (IP) injection of IHNV in juvenile herring with 10(3) pfu, IHNV was recovered from the tissues of sub-sampled individuals for only the first 5 d post-exposure. The lack of susceptibility to overt disease and transient levels of IHNV in the tissues of exposed fish indicate that Pacific herring do not likely serve a major epizootiological role in perpetuation of IHNV among free-ranging sockeye salmon Oncorhynchus nerka and farmed Atlantic salmon Salmo salar in the NE Pacific.     

Development and characterization of a cell line from Pacific herring, Clupea harengus pallasi, sensitive to both naphthalene cytotoxicity and infection by viral hemorrhagic septicemia virus

A cell line, PHL, has been successfully established from newly hatched herring larvae. The cells are maintained in growth medium consisting of Leibovitz's L-15 supplemented with 15% fetal bovine serum (FBS), and have been cryopreserved and maintain viability after thawing. These cells retain a diploid karotype after 65 population doublings. PHL are susceptible to infection by the North American strain of viral hemorrhagic septicemia (VHS) virus, and are sensitive to the cytotoxic effects of naphthalene, a common environmental contaminant. Naphthalene is a component of crude and refined oil, and may be found in the marine environment following acute events such as oil spills. In addition, chronic sources of naphthalene contamination include offshore drilling and petroleum contamination from areas such as docks and marinas that have creosote-treated docks and pilings and also receive constant small inputs of petroleum products. This cell line should be useful for investigations of the toxicity of naphthalene and other petroleum components to juvenile herring. In addition, studies of the VHS virus will be facilitated by the availability of a susceptible cell line from an alternative species.     

Biodiversity and exploitation of the main fish stocks in the Norwegian - Barents Sea ecosystem

Juvenile herring and capelin are the main stocks of plankton feeders in the Barents Sea, the cod is the dominant predator. Warm climate favours recruitment of herring and cod, but large stocks of juvenile herring hamper survival of the capelin fry. Since the early 1970s, the herring stock has been grossly overexploited, which could have led to an imbalance in the state of the predatorprey relationships in the Barents Sea. In the 1970s and early 80s, however, cod could feed on capelin which had excellent growth and recruitment conditions when the herring was absent. The consequences of the reduced herring stock were triggered in the mid 1980s, when excellent recruitment conditions for herring and cod occurred. Three abundant year classes of cod were recruited, but the herring stock was not sufficiently large to take full advantage of the favourable recruitment conditions. Given the lack of juvenile herring and a reduced capelin stock, the rapidly growing cod stock grazed down all other available prey species in the area, including its own progeny, and starved cod, seabirds and seals have in later years appeared on the north Norwegian coast. The capelin fishery collapsed, and the traditional coastal cod fisheries have been struck by the most serious crisis on record.     

Transmission dynamics of parasitic sea lice from farm to wild salmon

Marine salmon farming has been correlated with parasitic sea lice infestations and concurrent declines of wild salmonids. Here, we report a quantitative analysis of how a single salmon farm altered the natural transmission dynamics of sea lice to juvenile Pacific salmon. We studied infections of sea lice (Lepeophtheirus salmonis and Caligus clemensi) on juvenile pink salmon (Oncorhynchus gorbuscha) and chum salmon (Oncorhynchus keta) as they passed an isolated salmon farm during their seaward migration down two long and narrow corridors. Our calculations suggest the infection pressure imposed by the farm was four orders of magnitude greater than ambient levels, resulting in a maximum infection pressure near the farm that was 73 times greater than ambient levels and exceeded ambient levels for 30 km along the two wild salmon migration corridors. The farm-produced cohort of lice parasitizing the wild juvenile hosts reached reproductive maturity and produced a second generation of lice that re-infected the juvenile salmon. This raises the infection pressure from the farm by an additional order of magnitude, with a composite infection pressure that exceeds ambient levels for 75 km of the two migration routes. Amplified sea lice infestations due to salmon farms are a potential limiting factor to wild salmonid conservation.     

The life-cycle of the flatfish nematode Cucullanus heterochrous

Mature specimens of Cucullanus heterochrous Rudolphi, 1802 (Nematoda: Cucullanidae) were obtained from the intestine of the flounder, Platichthys flesus, from Danish waters. Eggs embryonate in seawater but do not hatch. Fully developed larvae pressed out of eggs are 430 microm long with amphids and dereids and enclosed within the cuticle of a previous larval stage. Infective larvae are believed to be in their third stage. Experimental studies showed that the polychaetes, Nereis spp., Scoloplos armiger, Brada villosa and Capitella sp., may act as intermediate hosts. In N. diversicolor the larvae increase their length to 1 mm within four weeks (15 degrees C) without moulting. Experimental infections showed that larvated eggs are not infective to fish, whereas >550 microm long larvae from polychaetes survived in 4-24 cm long flounders and plaice, Pleuronectes platessa. Third-stage larvae 550 microm to 1.1 mm long were found in the submucosa of the intestine one week post infection. At a length of about 800 microm to 1.4 mm they moult to fourth-stage larvae. Fourth-stage larvae, immature and mature worms occur in the intestine and rectum. Fourth-stage larvae and adults survived experimental transfer from one flounder to another. Similar developmental stages survived for two weeks in the intestine of experimentally infected cod, Gadus morhua.     

Ontogenic and ecological control of metamorphosis onset in a carapid fish, Carapus homei: experimental evidence from vertebra and otolith comparisons

In Carapus homei, reef colonisation is associated with a penetration inside a sea cucumber followed by heavy transformations during which the length of the fish is reduced by 60%. By comparing vertebral axis to otolith ontogenetic changes, this study aimed (i) to specify the events linked to metamorphosis, and (ii) to establish to what extent these fish have the ability to delay it. Different larvae of C. homei were caught when settling on the reef and kept in different experimental conditions for at least 7 days and up to 21 days: darkness or natural light conditions, presence of sea cucumber or not, and food deprivation or not. Whatever the nutritional condition, a period of darkness seems sufficient to initiate metamorphosis. Twenty-one days in natural light conditions delayed metamorphosis, whereas the whole metamorphosis process is the fastest (15 days) for larvae living in sea cucumbers. Whether the metamorphosis was initiated or not, otoliths were modified with the formation of a transition zone, whose structure varied depending on the experimental conditions. At day 21, larvae maintained in darkness had an otolith transition zone with more increments (around 80), albeit wider than those (more or less 21) of individuals kept under natural lighting. These differences in otolith growth could indicate an increased incorporation rate of released metabolites by metamorphosing larvae. However, the presence of a transition zone in delayed-metamorphosis larvae suggests that these otolith changes record the endogenously-induced onset of metamorphosis, whereas body transformations seem to be modulated by the environmental conditions of settlement.     

Manipulation of Developing Juvenile Structures in Purple Sea Urchins (Strongylocentrotus purpuratus) by Morpholino Injection into Late Stage Larvae

Sea urchins have been used as experimental organisms for developmental biology for over a century. Yet, as is the case for many other marine invertebrates, understanding the development of the juveniles and adults has lagged far behind that of their embryos and larvae. The reasons for this are, in large part, due to the difficulty of experimentally manipulating juvenile development. Here we develop and validate a technique for injecting compounds into juvenile rudiments of the purple sea urchin, Strongylocentrotus purpuratus. We first document the distribution of rhodaminated dextran injected into different compartments of the juvenile rudiment of sea urchin larvae. Then, to test the potential of this technique to manipulate development, we injected Vivo-Morpholinos (vMOs) designed to knock down p58b and p16, two proteins involved in the elongation of S. purpuratus larval skeleton. Rudiments injected with these vMOs showed a delay in the growth of some juvenile skeletal elements relative to controls. These data provide the first evidence that vMOs, which are designed to cross cell membranes, can be used to transiently manipulate gene function in later developmental stages in sea urchins. We therefore propose that injection of vMOs into juvenile rudiments, as shown here, is a viable approach to testing hypotheses about gene function during development, including metamorphosis.     

The effects of thin layers on the vertical distribution of larval Pacific herring, Clupea pallasi

Temporal and spatial heterogeneity of resources is likely to have dramatic effects on the behaviors of zooplankton and ichthyoplankton, which in turn are likely to have strong effects on ecological dynamics such as predation, growth, and mating. The objective of this study was to determine whether vertically thin layers of extreme prey concentration affect the vertical distribution of larval Pacific herring (Clupea pallasi). We employed 2-m tall experimental tanks equipped with video cameras that scanned the vertical extent of the tanks to investigate the effects of thin layers on the vertical distribution of 5- and 10-day-old herring larvae. Three treatments were established: (1) a thin layer of prey (rotifers, Brachionus plicatilis) through density (salinity) stratification, (2) homogeneous vertical distribution of both prey and density, and (3) density (salinity) stratification, but with a homogeneous distribution of prey. We found that in all treatments the majority of larval herring were at the surface, near the light, despite the absence of a peak in rotifer abundance at this depth in some instances. However, there were also clear effects of the thin layers—secondary subsurface peaks in herring abundance occurred at the mid-depths in the stratified tanks, in and around the thin layers. In addition, our results provide some evidence that thin layers specifically, rather than prey patches generally, influence the vertical distribution of larval herring, i.e., larvae may use the physical properties of thin layers to locate and distribute themselves, instead of reacting solely to the prey patches. Thus thin layers can affect directly the vertical distribution of larval herring, and perhaps indirectly their horizontal distribution, as herring larvae live in environments (e.g., estuaries) where advective transport is also often vertically heterogeneous.     

Validation of daily increment formation in otoliths for three Diplodus species in the Mediterranean sea

The efficacy of two fluorochromes, chlorhydrate of tetracycline (CHTC) and alizarin complexone (ALC), to induce a label on the otoliths of juvenile sparid fishes by immersion techniques was tested in different experimental conditions. CHTC did not mark otoliths in natural sea water and was toxic in divalent cation-free sea water. Immersing fish in a 50-mg 1⁻¹ ALC natural seawater solution for 24 h induced a well-defined mark on the otoliths and had no effect on survival. A daily periodicity of increment formation on otoliths was observed in captivity for Diplodus vulgaris and D. puntazzo , and was conserved in natural environment for D. sargus . The frequency of increment deposition did not vary with the age of juvenile fishes. Thus, otolith microstructure analysis will be a reliable method to give age estimates in these three sparid species during their juvenile life.     

Effect of larval ontogeny, turbulence and light on prey attack rate and swimming activity in herring larvae

The effects of ontogeny (larval size), light and turbulence on the attack rate and swimming activity (proportion of time swimming and duration of swimming bout) of herring larvae (15-28 mm TL) have been investigated. Emphasis was put on the experimental design in order to create a set-up where the turbulence intensity distribution could be accurately measured as well as controlled in the entire experimental tank.Both larval size (ontogeny) and light had a significant positive effect on prey attack rate. Likewise, an intermediate increase in turbulence had a positive effect on prey attack rate, but this effect was dependent of light intensity and larval size.At low light (1.5 μE m² s⁻¹) intermediate turbulence increased the prey attack rate significantly for larger larvae (26 and 28 mm), while at high light (18 μE m² s⁻¹) intermediate turbulence had only a significant positive effect on the attack rate of smaller larvae 20 and 23 mm.In general, our data show a dome-shaped response of turbulence on attack rate and a U-shaped response of turbulence on swimming activity.For herring larvae >20 mm, the maximum (attack rate) and minimum (swimming activity) response of turbulence were found at intermediate turbulence intensities (energy dissipation rates between 7∗10⁻⁸ and 1∗10⁻⁶ W/kg). The highest turbulence level tested (8∗10⁻⁶ W/kg) showed only negative effects, as attack rates where at the lowest and swimming activity at the highest.Swimming activity increased with larval size or light, and decreased at intermediate turbulence. Compared to turbulent intensities under natural conditions this implies that larger herring larvae at 10 m depth have to be exposed to wind speeds of more than 17 m/s before negative effects on attack rate and swimming activity occurs.     

The development of schooling behaviour in Atlantic herring Clupea harengus

The timing of the development of schooling behaviour in Atlantic herring Clupea harengus and the quantitative and qualitative evolution of such behaviour in the larval and post-metamorphic stages were investigated with a simulation model and experimental observations of laboratory-reared fish. Herring larvae started to develop schooling behaviour between the lengths of 35 and 40 mm. This coincided with the beginning of metamorphosis, characterized by ontogenetic changes which require a switch to a different antipredator strategy from that employed by early larvae, while providing the sensory and anatomical mechanisms necessary for the formation of schools. Schooling behaviour was established at the end of metamorphosis (50–55 mm) and its characteristics continued to evolve throughout the early juvenile stage. A critical period in terms of vulnerability to predation is expected between the end of the larval and beginning of the juvenile stages, before schooling behaviour is fully developed. The implications of some of the results for the methodology of future studies are also discussed.     

Parental effects on size variation in fish larvae

The growth and changes in the standard deviation about mean length were measured for sole, Solea solea L., and herring, Clupea harengus L., larvae from crosses of one or two males with one or two females. The sizes of sole larvae did not differ between crosses but the size variation increased faster with age when two females were used. Both maternal and paternal effects were detected for size at hatching in herring, but there was no parental influence on size variation. Size variation in herring larvae declined or was stable during the yolk sac stage and was not affected by the number of parents used for fertilizations.     

Ecology of larval herring in relation to the oceanography of the Gulf of Maine

The dynamics of larval herring in the north-eastern Gulf ofMaine are reviewed with reference to the important physicaland biological oceanographic processes of the region. Particularattention is given to the apparently conflicting hypothesesof larval drift from the tidally well-mixed spawning areas tonursery areas, and larval retention in spawning areas for aperiod of several months. Both processes have been reportedfor the eastern Maine-Grand Manan spawning area of the Gulfof Maine, but the relative importance of each to larval survivalthrough the winter and recruitment to the juvenile stage isnot clear. Both transport and retention are interpreted in lightof oceanographic processes that might impart variability, especiallybetween years, in the proportion of larvae transported awayversus that retained. Results of recent survey cruises in theGulf of Maine show both hypotheses to have merit. It is suggestedthat (i) interannual differences in slope water intrusions intothe Gulf of Maine as they affect the residual circulation, (ii)the lunar periodicity in the intensity of tidal mixing in relationto hatching times, and (iii) the potential for variable spawninglocations relative to the tidal fronts, may affect the distributionsof herring larvae immediately after hatching in the fall, andmay control the proportion of larvae that are advected awaywith the residual currents versus that retained in the vicinityof spawning. It is also suggested that those processes thataffect larval distributions and survival in the fall are importantin determining the overwintering distributions of larvae inthe Gulf of Maine, where the subtle influences of variable foodregimes and water temperatures could potentially exact largeinterannual differences in winter survival and recruitment tothe juvenile stage.     

Ultrastructural changes in the muscle tissue of Clupea harengus L. larvae induced by acid pH

The pH of sea water remains relatively constant at 7·8–8·3 but, in lower salinity environments, inshore and in estuaries, localized changes in pH can be caused by effluent discharge. Eggs and larvae of the Atlantic herring, Clupea harengus L., can be spawned and develop in these areas and thus may be exposed to acidic conditions. Larvae hatched from eggs exposed to sea water at pH 7·0 or 6·0 or 5·2, in comparison to controls, showed a significant reduction in the mean volume of muscle fibres and mitochondria, degeneration of mitochondrial cristae and swelling of the sarcoplasmic reticulum. These sublethal ultrastructural changes would cause reduced metabolic activity and loss of contractility leading to reduced swimming and equilibrium ability with consequent loss of viability.