Inheritance of allozymes in Atlantic herring (Clupea harengus harengus)

Progeny from single pair crosses of Atlantic herring were examined to determine the heritability of genetic variation at seven polymorphic allozyme loci. Mendelian inheritance of codominant autosomal alleles was established for IDH-2, LDH-1, LDH-2, ME-2, PGM-1, and PGI-2. This demonstration of Mendelian inheritance is essential for accurate interpretation of allozymic variation among natural populations of this pelagic species.     

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.     

Fourteen new microsatellite markers for Atlantic herring Clupea harengus

Fourteen new microsatellite loci were developed and tested on Atlantic herring Clupea harengus with 39 individuals from Iceland and 49 individuals from Norway. The microsatellites, which contain di, tri and tetranucleotide repeats, are polymorphic (7-30 alleles), with observed heterozygosity ranging between 0·69 and 1·00 and expected heterozygosity between 0·55 and 0·97.     

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.     

Cholecystokinin (CCK) in Atlantic herring (Clupea harengus L.) - ontogeny and effects of feeding and diurnal rhythms

Neural and alimentary cholecystokinin (CCK) levels in Atlantic herring, Clupea harengus, were analyzed from hatching to 40days after hatching (DAH). The head compartment representing the neural pool was quantitatively dominant (>80% of the total CCK content) while the digestive tract pool represented 6-10%. During ontogeny the CCK level in whole larvae increased almost 15-fold from 0 to 40 DAH, being particularly marked from 14 to 20 DAH. Larvae of 24 to 26 DAH were examined for potential occurrence of a circadian rhythm and to analyze the effects of feeding. Fed and fasted larvae were significantly different, where fed larvae showed higher CCK levels. There were large fluctuations in CCK levels analyzed at 3h intervals without an apparent diurnal pattern. Shorter sampling intervals of 1h in the morning when lights were switched gradually on and food was offered to the larvae demonstrated a marked drop in the relative gut CCK levels and a concurrent increase in the CCK carcass to gut ratio, 1h after introduction of food followed by a return to prefeeding levels after 2h. This response probably results from a release and re-synthesis of CCK in the gut after initiation of feeding. Taken together, these results support earlier reports that CCK participates in the regulation of digestive processes in herring larvae, but CCK does not seem to have a circadian rhythm independent of feeding.     

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.     

Zuchtversuche mit Heringslarven (Clupea harengus L.)

Ohne ZusammenfassungMit 4 Abbildungen im Text.Anm. Die Arbeit wurde durch eine Beihilfe der deutschen Forschungsgemeinschaft ermöglicht.     

Variation in egg mass within two Atlantic herring Clupea harengus stocks

The main objective of this study was to investigate if egg size (mass) at spawning is invariant for Scotia-Fundy summer and autumn (SFSH) and Icelandic summer (ISSH) spawning herring Clupea harengus. Oocyte dry mass measurements for SFSH females collected in 2001 and ISSH females collected in 1999 and 2000 showed a large variation. Difference in egg dry mass among fish was found to vary by as much as twofold in each stock. For ISSH, variation in egg mass was also apparent from oocyte volume measurements made jointly with a histological examination of the ovaries. Approximately 20% of the variation in egg mass could be explained by maternal whole-body mass or total length, indicating that length or age composition in the stocks can potentially influence the recruitment success. This implies that fisheries management strategies should aim to maintain a broad range in age composition.     

Otolith shape: a population marker for Atlantic herring Clupea harengus

Otolith shape variation of seven Atlantic herring Clupea harengus populations from Canada, the Faroe Islands, Iceland, Ireland, Norway and Scotland, U.K., covering a large area of the species' distribution, was studied in order to see if otolith shape can be used to discriminate between populations. The otolith shape was obtained using quantitative shape analysis, transformed with Wavelet and analysed with multivariate methods. Significant differences were detected among the seven populations, which could be traced to three morphological structures in the otoliths. The differentiation in otolith shape between populations was not only correlated with their spawning time, indicating a strong environmental effect, but could also be due to differing life‐history strategies. A model based on the shape differences discriminates with 94% accuracy between Icelandic summer spawners and Norwegian spring spawners, which are known to mix at feeding grounds. This study shows that otolith shape could become an accurate marker for C. harengus population discrimination.     

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.     

Morphological and meristic differences among spawning aggregations of north-east Atlantic herring, Clupea harengus L.

Univariate and multivariate analyses were performed on seven anatomical characters of Atlantic herring, Clupea harengus , sampled from seven discrete spawning grounds to the west of the British Isles and from spawning sites in the Baltic and Thames Estuary. Analysis of mean similarities, Mahalanobis D², percentage overlap and projections from prinicipal co-ordinate analysis demonstrated that a high degree of anatomical similarity exists among the herring spawning communities to the west of the British Isles. Although local aggregations had distinct centres of abundance they appeared to have large overlapping distributions. However, the Celtic Sea sample was found to be distinctly different from adjacent and neighbouring stocks. The samples of coastal herring from the Thames Estuary and the Baltic Sea showed little overlap with other samples.     

Zum Formwachstum des herings (Clupea harengus L.)

Zusammenfassung Die Herausbildung der Körperform und die Umwachsung der Leibeshöhle wird an Hand von Querschnitten hinter der Dorsalflosse verfolgt. Dazu wurden 43 Heringe von 5–245 mm Körperlänge untersucht und mit dem Planimeter vermessen.Während der relative Eingeweidequerschnitt zeitlebends ziemlich konstant bleibt, abgesehen von frühesten Stadien und laichreifen Tieren, wächst der Stützgewebequerschnitt negativ, der Muskelquerschnitt positiv allometrisch bis zur Metamorphose, um dann annähernd isometrisch zuzunehmen.Die Seitenmuskulatur umwächst, am Schultergürtel beginnend, die Leibeshöhle. Dadurch ergeben sich für jeden Querschnittsort eigene Daten, wann der Zustand des ausgeformten Tieres erreicht wird. Für den hier untersuchten Querschnitt hinter der Dorsalen fällt dieser Zeitpunkt in den Beginn der Metamorphose.Der Umfang erreicht ebenfalls in der Metamorphose die relativen Werte des ausgeformten Tieres, er nimmt, wie zu erwarten, bei der Reifung zu. Die fur die fertigen Tiere charakteristische Umrißlinie wird schon bei der 15 mm langen Larve ausgebildet, durch die Gonadenreifung tritt wiederum eine Annäherung an die Kreisform ein.     

Marine landscapes and population genetic structure of herring (Clupea harengus L.) in the Baltic Sea

Numerically small but statistically significant genetic differentiation has been found in many marine fish species despite very large census population sizes and absence of obvious barriers to migrating individuals. Analyses of morphological traits have previously identified local spawning groups of herring (Clupea harengus L.) in the environmentally heterogeneous Baltic Sea, whereas allozyme markers have not revealed differentiation. We analysed variation at nine microsatellite loci in 24 samples of spring-spawning herring collected at 11 spawning locations throughout the Baltic Sea. Significant temporal differentiation was observed at two locations, which we ascribe to sympatrically spawning but genetically divergent 'spawning waves'. Significant differentiation was also present on a geographical scale, though pairwise F(ST) values were generally low, not exceeding 0.027. Partial Mantel tests showed no isolation by geographical distance, but significant associations were observed between genetic differentiation and environmental parameters (salinity and surface temperature) (0.001 < P < or = 0.099), though these outcomes were driven mainly by populations in the southwestern Baltic Sea, which also exhibits the steepest environmental gradients. Application of a novel method for detecting barriers to gene flow by combining geographical coordinates and genetic differentiation allowed us to identify two zones of lowered gene flow. These zones were concordant with the separation of the Baltic Sea into major basins, with environmental gradients and with differences in migration behaviour. We suggest that similar use of landscape genetics approaches may increase the understanding of the biological significance of genetic differentiation in other marine fishes.     

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.     

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.