Variation in body condition, fat content and growth rate of Icelandic summer-spawning herring Clupea harengus L.

Interannual variation in Fulton's condition factor ( K ), total body-fat content and growth rate were investigated for the Icelandic summer-spawning herring Clupea harengus over the period from 1963 to 1999. The results show a close relationship between K and the body-fat content in the autumn, both of which correlated to the growth rate in the same calendar year. A relationship was also found between K during the prespawning season (February to June) and K in the autumn (September to December) in the same year. The different measures of body condition were not found to be related to sea temperature or zooplankton indices, while the spawning stock biomass had negative effects on the fish condition. The results imply that the body condition in the autumn is representative of the feeding success over the whole foregoing year and is therefore related to the growth rate that year. The growth rate appears to be relatively uniform throughout the year.     

Identification of male heterogametic sex-determining regions on the Atlantic herring Clupea harengus genome

The sex determination system of Atlantic herring Clupea harengus L., a commercially important fish, was investigated. Low coverage whole-genome sequencing of 48 females and 55 males and a genome-wide association study revealed two regions on chromosomes 8 and 21 associated with sex. The genotyping data of the single nucleotide polymorphisms associated with sex showed that 99.4% of the available female genotypes were homozygous, whereas 68.6% of the available male genotypes were heterozygous. This is close to the theoretical expectation of homo/heterozygous distribution at low sequencing coverage when the males are factually heterozygous. This suggested a male heterogametic sex determination system in C. harengus, consistent with other species within the Clupeiformes group. There were 76 protein coding genes on the sex regions but none of these genes were previously reported master sex regulation genes, or obviously related to sex determination. However, many of these genes are expressed in testis or ovary in other species, but the exact genes controlling sex determination in C. harengus could not be identified.     

Influence of temperature on evasive responses of Atlantic herring larvae attacked by yearling herring, Clupea harengus L.

Predation encounters were staged in the laboratory between yearling herring, Clupea harengus L., 66 to 104 mm t.l. , and herring larvae, 8 to 30 mm t.l. ., at 8,11 and 14 o C. Video records were used to quantify prey behaviour. Prey responsiveness, reactive distance, response latency, and apparent looming threshold were not affected by temperature. Response speeds increased with temperature. Predator error rate and capture success showed no consistent thermal effects. Although the experiments could not fully evaluate the influence of temperature on the predators, results suggest that the predator's performance largely governs the outcome of an attack on a larva and that higher temperatures favour the predator by increasing the frequency of its encounter with prey.     

Routine respiration rates of Atlantic mackerel, Scomber scombrus L., and herring, Clupea harengus L., at low activity levels

Once adapted to the captive environment, mean minimum respiration rates were 118 mgO₂ kg⁻¹ h⁻¹ for mackerel, body length ( b.l ) range 290 to 380 mm, at 11.1o C at a swimming speed of 0.6 b.l. s¹ and 93 mgO₂ kg⁻¹ h¹ for herring, length range 255 to 310 mm, at 9.3° C at a swimming speed of 0.3 b.l. s¹.     

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.     

Population structure of Atlantic herring, Clupea harengus L.

During the last decade, genetic studies have been carried out on samples of Atlantic herring, CIupea harengus L., from the Baltic, North Sea, British Isles, Norwegian sea-waters and in the western part of the North Atlantic Ocean. Based on direct comparisons of samples drawn from several areas, the available information on allozyme variation was compiled into a comparable data set and analysed for population structure and genetic distance. The results are discussed in relation to the present stock model for herring.     

The relative profitability of particulate- and filter-feeding in the herring, Clupea harengus L.

Analysis of laboratory video-recordings of herring feeding by biting and filtering on Calanus finmarchicus and three sizes of Artemia enabled the capture rates of the two methods to be estimated at different prey concentrations. At low concentrations the fish feed by selective capture of individual particles, but the capture rate achievable by this method is constrained by the maximum rate at which they can bite. Filter-feeding is not subject to this constraint because capture rate is directly proportional to prey concentration and above a critical prey concentration its capture rate exceeds that of biting. The possession of two feeding methods allows the fish to maximize prey intake over a wide range of concentrations and the phenomenon of switching between feeding methods can be explained by their relative profitabilities at different concentrations. The observation that less than 50% of fish are filtering when capture rates by the two methods are equal suggests that filtering is energetically more costly than biting. Estimates of the energy cost of filtering indicated that it may be from 1.4 to 4.6 times higher than that of biting.     

Differential utilization of storage lipids and storage proteins by Northwest Atlantic herring (Clupea harengus harengus)

Utilization of storage lipids and storage proteins by non-feeding north-west Atlantic herring was assessed under conditions of: (1) concurrent gonad maturation and overwintering periods with differing rates of gonad development for males and females (spring-spawncrs); (2) overwintering, reproductively dormant and spent males and females (autumn-spawners); and (3) approximately equivalent rates of gonad development during late summer for males and females (autumn-spawners). Storage lipids were depleted by all non-feeding fish but storage proteins were depleted only by herring with measurable gonad development during the observation period independently of season (female spring-spawners, maturing male and female autumn-spawners). The analyses indicate that storage lipids sustain routine and active metabolism while storage proteins are utilized to develop gonad. Therefore, the relative investment into reproduction versus routine and active metabolism can be evaluated separately for non-feeding herring using readily obtainable tissue proximate composition data.     

Characterization of a major histocompatibility class II A gene (Clha-DAA) with an embedded microsatellite marker in Atlantic herring (Clupea harengus L.)

An Atlantic herring major histocompatibility class II A ( Clha-DAA ) cDNA sequence has been characterized and was shown to encode a leader peptide, alpha-1 domain, alpha-2 domain, connecting peptide, transmembrane and cytoplasmic region. The Clha-DAA protein sequence has all the characteristics of a teleost class II A protein with conserved cysteines in both the alpha-1 and the alpha-2 domains and two potential N-linked glycosylation sites. Exon 2 sequences encoding the polymorphic alpha-1 domain from different individuals were analysed and revealed the presence of at least two loci. The Clha-DAA gene consists of four exons and three short introns. Four unique intron 3 sequences from multiple individuals were obtained and were shown to contain a (TG)_n microsatellite sequence. Primers were optimized such that only a single microsatellite locus designated Clha-DAA-INTR3 was amplified. Four herring populations from the North Sea and the Baltic Sea were genotyped for Clha-DAA-INTR3 . In total, 16 Clha-DAA-INTR3 alleles were detected; the distribution of the alleles showed no deviation from Hardy–Weinberg expectation. Levels of genetic differentiation among samples were of similar magnitude as have been reported earlier for neutral microsatellite loci between northern North Sea and Baltic Sea herring populations.     

Comparative study of early-life survival variability of herring Clupea harengus, in the north-eastern Atlantic

Variation in the survival of herring between the egg stage and the age of recruitment to the spawning population has been examined for seven Atlantic herring populations in northern European waters. The sources of the variation have been partitioned between density-dependent and density-independent factors. The magnitude of the density-independent component was found to be related to the scale of the processes controlling the dispersion and distribution of larvae from the various populations. We conclude that the spawning strategies of some populations are adapted to oceanographic conditions to maximize the probability of a particular larval transport pattern, although no single model could account for all spawning strategies.     

Development of microsatellite loci in Pacific herring (Clupea pallasi)

Fifteen polymorphic di‐, tri‐ and tetranucleotide microsatellite markers were developed in Pacific herring (Clupea pallasi) to aid in the delineation of population structure of herring in British Columbia. Twelve of the microsatellite loci were analysed in over 4000 wild herring. Expected heterozygosities ranged 0.73–0.95, and only two loci significantly deviated from Hardy–Weinberg equilibrium.     

The effect of progressive hypoxia on swimming activity and schooling in Atlantic herring

Schools of herring exposed to progressive hypoxia show a peak in velocity during severe hypoxia, at 15–34% oxygen saturation, followed by a decrease in swimming speed until school disruption occurred. The observed increase in swimming speed during severe hypoxia reveals a graded response, since the lower the fish's swimming speed prior to severe hypoxia ( U ₉₅₋₅₀, the speed at oxygen saturations between 95 and 50%), the greater the relative increase in swimming speed. The oxygen saturations at which both peak velocity and school disruption occurred were lower for fish with lowest U ₉₅₋₅₀, suggesting that the fish with the slowest speed U ₉₅₋₅₀ reach their critical P_O2 (at which there is respiratory distress) last, i.e. at lower oxygen saturation. At a functional level, it is suggested that herring encountering hypoxia increase their speed in order to find more favourable conditions, and the magnitude of this increase is modulated by their respiratory distress. It is also hypothesised that the observed increase in speed may be related to an increase in the rate of position shifting within the school. Since the oxygen saturation at which the response to hypoxia occurs and the magnitude of the response are related to the fish's preferred speed prior to severe hypoxia, it is suggested that such a preferred speed should be measured in experiments testing the effect of hypoxia on fish behaviour.     

Length of herring larvae in relation to age and time of hatching

The lengths of newly hatched herring Clupea harengus larvae were generally larger than the lengths of embryos released from eggs on the same day. Over subsequent days, the daily increase in length was higher in groups that were released from the eggs as opposed to groups that hatched naturally. Thus by the end of the hatching period (18 days after fertilization) there were no significant differences in mean lengths.     

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.     

Effects of two Avtemia diets with different contents of polyunsaturated fatty acids on the lipid composition of larvae of Atlantic herring (Clupea harengus

Atlantic herring larvae ( Clupea harengus ) were fed two enriched Artemia diets with different contents of (n-3) highly unsaturated fatty acids (HUFA), one containing low levels of 20: 5(n-3) and no 22: 6(n-3), the other containing substantial levels of both 20: 5(n-3) and 22: 6(n-3). After 30 days of culture, fatty acid compositions of lipid classes in the heads, bodies and eyes of the larvae were analysed. Fish fed Artemia with the low (n-3) HUFA diet lacking 22: 6(n-3) had lower amounts of total (n-3)HUFA and, in particular, of 22:6(n-3) in individual phospholipids and total neutral lipids of heads, bodies and eyes as compared to fish fed Artemia with high levels of (n-3)HUFA. The amount of 22: 6(n-3) in the fatty acids of phosphatidyl-ethanolamine of eyes was particularly susceptible to dietary depletion. The implications of these findings are discussed, particularly in relation to dietary requirements for 22: 6(n-3) during development of neural tissue in predatory fish iarvae.     

Duration of synchronous cleavage cycles and rate of development at different temperatures in the Baltic herring

The duration of one synchronous cleavage cycle (τ₀) in Clupea harengus membras at different temperatures ( T ) was given by: (logτ₀)= 2.4349–0–0684T for T= 0.9–13°C, and (logτ₀)= 1.61010–oooit for t= 13–18–7°C.     

Environmental correlates of population differentiation in Atlantic herring

The marine environment is characterized by few physical barriers, and pelagic fishes commonly show high migratory potential and low, albeit in some cases statistically significant, levels of genetic divergence in neutral genetic marker analyses. However, it is not clear whether low levels of differentiation reflect spatially separated populations experiencing gene flow or shallow population histories coupled with limited random genetic drift in large, demographically isolated populations undergoing independent evolutionary processes. Using information for nine microsatellite loci in a total of 1951 fish, we analyzed genetic differentiation among Atlantic herring from eleven spawning locations distributed along a longitudinal gradient from the North Sea to the Western Baltic. Overall genetic differentiation was low (theta = 0.008) but statistically significant. The area is characterized by a dramatic shift in hydrography from the highly saline and temperature stable North Sea to the brackish Baltic Sea, where temperatures show high annual variation. We used two different methods, a novel computational geometric approach and partial Mantel correlation analysis coupled with detailed environmental information from spawning locations to show that patterns of reproductive isolation covaried with salinity differences among spawning locations, independent of their geographical distance. We show that reproductive isolation can be maintained in marine fish populations exhibiting substantial mixing during larval and adult life stages. Analyses incorporating genetic, spatial, and environmental parameters indicated that isolating mechanisms are associated with the specific salinity conditions on spawning locations.     

Vulnerability of late larval and early juvenile Atlantic herring, Clupea harengus, to predation by whiting,Merlangius merlangus

Laboratory experiments investigated changes in the vulnerability of herring to predation by whiting during growth from <34 to >80mm in length, Premetamorphic herring (<50mm) failed to react to c . 50% of attacks by the predator, but this was reduced to c . 20% in postmetamorphic fish. Premetamorphic herring failing to react had c. 30% probability of survival due to unforced errors by the predator (unsuccessful attacks that did not elicit a reaction by the prey), but these errors did not occur after herring metamorphosis. The loss of the advantage of lower conspicuousness following metamorphosis was counterbalanced by increased reactivity. Antipredator benefits should increase with schooling behaviour (first evident in experiments at c . 50 mm), so predation mortality should decrease in postmetamorphic herring in the sea. Metamorphosis itself would be associated with high predation risk since conspicuousness is enhanced but reactivity and schooling behaviour are still not fully developed.     

Effect of spatial differences in growth on distribution of seasonally co‐occurring herring Clupea harengus stocks

The mechanisms most likely to determine the distribution of the two major herring Clupea harengus stocks in their common early summer feeding ground in the eastern North Sea, Skagerrak and Kattegat were investigated through analysis of acoustic survey data from six consecutive years. No change was detected in biomass of North Sea autumn spawning C. harengus (NSAS) over time, whereas the biomass of western Baltic spring spawning C. harengus (WBSS) declined severely. Analyses of centre of abundance by stock showed no change in NSAS distribution, whereas the WBSS changed to a more western distribution over time. Contrary to previous perception of the juvenile migration, NSAS were found to leave the study area at the age between 1 and 2 years and WBSS 1 year olds were encountered in the Skagerrak. The estimated parameters of von Bertalanffy growth equations showed marked differences between areas with fish in the eastern part of the area having the lowest size at age at all ages. Further, their growth conditions appeared to deteriorate progressively over the period studied. Both NSAS and WBSS showed the highest condition in the North Sea and Skagerrak while condition was substantially lower in age Kattegat. The westward movement of spring spawners over time suggests that growth rate and possibly density of conspecifics influence the migration pattern and distribution of C. harengus in the area. In contrast, there was no evidence to suggest that distribution was constant over time within stocks or that distribution reflected size‐dependent limitations on migration distance.     

Differences in temperature conditions and somatic growth rate of larval and early juvenile spring-spawned herring from the Vistula Lagoon, Baltic Sea manifested in the otolith to fish size relationship

Larval and juvenile herring Clupea harengus collected in the Polish part of the Vistula Lagoon in May-July 1997 had hatched between 17 April and 9 June and originated from three cohorts. The spawning season began on 1 March at 3·8° C and was completed on 3 June at 12·7° C. Mortality among larvae was high in the first 2 weeks of April, probably associated with significant temperature decrease at the beginning of the spawning season. The growth of 10–48 mm L _S herring was linear, highest for larvae and juveniles from the first cohort (0·58 mm mm^-1 day^-1), slower for the second cohort (0·55 mm mm^-1 day^-1) and the slowest for the third cohort (0·45 mm mm day^-1). Temperature effects on the growth were inconclusive and potentially unfavourable feeding conditions in June might have been responsible for the relatively slow growth of third cohort larvae and juveniles.
Relationships between otolith size (perimeter, length, width, area, and weight) and fish size ( L _S) differed among the three cohorts, related mostly to the positive temperature effect on otolith growth, individuals growing in warmer water had larger otoliths. Although a negative growth rate effect was observed as well, it was less significant.