Seasonal Dynamics of Atlantic Herring (Clupea harengus L.) Populations Spawning in the Vicinity of Marginal Habitats

Gillnet sampling and analyses of otolith shape, vertebral count and growth indicated the presence of three putative Atlantic herring (Clupea harengus L.) populations mixing together over the spawning season February–June inside and outside an inland brackish water lake (Landvikvannet) in southern Norway. Peak spawning of oceanic Norwegian spring spawners and coastal Skagerrak spring spawners occurred in March–April with small proportions of spawners entering the lake. In comparison, spawning of Landvik herring peaked in May–June with high proportions found inside the lake, which could be explained by local adaptations to the environmental conditions and seasonal changes of this marginal habitat. The 1.85 km² lake was characterized by oxygen depletion occurring between 2.5 and 5 m depth between March and June. This was followed by changes in salinity from 1–7‰ in the 0–1 m surface layer to levels of 20–25‰ deeper than 10 m. In comparison, outside the 3 km long narrow channel connecting the lake with the neighboring fjord, no anoxic conditions were found. Here salinity in the surface layer increased over the season from 10 to 25‰, whereas deeper than 5 m it was stable at around 35‰. Temperature at 0–5 m depth increased significantly over the season in both habitats, from 7 to 14°C outside and 5 to 17°C inside the lake. Despite differences in peak spawning and utilization of the lake habitat between the three putative populations, there was an apparent temporal and spatial overlap in spawning stages suggesting potential interbreeding in accordance with the metapopulation concept.     

Zuchtversuche mit Heringslarven (Clupea harengus L.)

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

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.     

Latitudinal Gradient in Otolith Shape among Local Populations of Atlantic Herring (Clupea harengus L.) in Norway

Otolith shape analysis of Atlantic herring (Clupea harengus) in Norwegian waters shows significant differentiation among fjords and a latitudinal gradient along the coast where neighbouring populations are more similar to each other than to those sampled at larger distances. The otolith shape was obtained using quantitative shape analysis, the outlines were transformed with Wavelet and analysed with multivariate methods. The observed morphological differences are likely to reflect environmental differences but indicate low dispersal among the local herring populations. Otolith shape variation suggests also limited exchange between the local populations and their oceanic counterparts, which could be due to differences in spawning behaviour. Herring from the most northerly location (69°N) in Balsfjord, which is genetically more similar to Pacific herring (Clupea pallasii), differed in otolith shape from all the other populations. Our results suggest that the semi-enclosed systems, where the local populations live and breed, are efficient barriers for dispersal. Otolith shape can thus serve as a marker to identify the origin of herring along the coast of Norway.     

Detection of small bioactive peptides from Atlantic herring (Clupea harengus L.)

Recent research has shown that fish residual materials contain a range of components with interesting biological activity. Therefore, there is a great potential in the marine bioprocess industry to utilize these by-products as starting material for generating more valuable products. The aim of the present study was to search for bioactive peptides (in particular small natural bioactive peptides with molecular weight lower than 10 kDa) in Atlantic herring (Clupea harengus L.) by-products such as skin and more general residual materials. By such means a range of peptides with claimed interesting biological activities was found. Herein the activity of the detected bioactive peptides and strategies for isolating peptide fragments containing the bioactive motif is discussed. Identification of bioactive peptides in crude peptide/protein sources (skin and residual materials) was performed directly using a combination of mass spectrometry (Orbitrap), bioinformatics and database search. This method was a good angle of approach in order to map the potential in new species and species that have been very little studied.     

Direct Effects of Microalgae and Protists on Herring (Clupea harengus) Yolk Sac Larvae

This study investigated effects of microalgae (Rhodomonas baltica) and heterotrophic protists (Oxyrrhis marina) on the daily growth, activity, condition and feeding success of Atlantic herring (Clupea harengus) larvae from hatch, through the end of the endogenous (yolk sac) period. Yolk sac larvae were reared in the presence and absence of microplankton and, each day, groups of larvae were provided access to copepods. Larvae reared with microalgae and protists exhibited precocious (2 days earlier) and ≥ 60% increased feeding incidence on copepods compared to larvae reared in only seawater (SW). In the absence and presence of microalgae and protists, life span and growth trajectories of yolk sac larvae were similar and digestive enzyme activity (trypsin) and nutritional condition (RNA-DNA ratio) markedly declined in all larvae directly after yolk sac depletion. Thus, microplankton promoted early feeding but was not sufficient to alter life span and growth during the yolk sac phase. Given the importance of early feeding, field programs should place greater emphasis on the protozooplankton-ichthyoplankton link to better understand match-mismatch dynamics and bottom-up drivers of year class success in marine fish.     

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.     

Annual survival of adult Atlantic Puffins Fratercula arctica is positively correlated with Herring Clupea harengus availability

Atlantic Herring is a keystone species in several marine ecosystems, supporting intensive fisheries as well as many predators including seabirds. Biomass of this stock in eastern North America has declined considerably in recent years, potentially putting at risk populations of its predators. Although adult survival in seabirds is considered robust to moderate changes in food availability, it is also the life‐history component most critical to sustaining populations of long‐lived birds. To investigate the possibility that Atlantic Puffin survival has been affected by reduced abundance of its main prey, we analysed the encounter histories of 2999 Atlantic Puffins ringed on Machias Seal Island to estimate annual adult survival for the years 1999–2011 and assess trends in survival and the effects of several biological and environmental covariates. Features of Puffin biology and resighting procedures likely to introduce heterogeneity into our resighting probabilities were accounted for and models of survival were assessed using standard methods. We used the variance components procedure in Program MARK and survival estimates from a time‐varying model to estimate the process variance (biological variation in survival) accounted for by suspected covariates of survival. Two proxies of food availability each explained more than half of the variation in annual survival: fishery landings of Atlantic Herring (52%) and per cent (by mass) of 1‐group Herring in the diet of Puffin chicks (51%). In addition to these proxies, May sea‐surface temperature accounted for 37% of variance in survival, but winter values of North Atlantic Oscillation showed no effect. Of those parameters of Puffin biology examined, chick growth rate explained 19% of the process variance in annual survival; laying date, fledging condition and fledging date all explained no variance. A decline in fishery landings of Herring since the early 1990s, and a concurrent decline in adult Puffin survival, reinforces concern for the health of the population of Herring, a keystone forage fish in this region, and of the community of marine predators in the Gulf of Maine that rely on Herring for their survival and reproduction.     

Separating two herring stocks from growth data: long‐term changes in survey indices for Western Baltic Spring Spawning Herring (Clupea harengus) after application of a stock separation function

In the Baltic Sea, herring stocks are surveyed and managed according to a spatial allocation based on ICES (International Council for the Exploration of the Sea) subdivisions. In the western Baltic, the distribution areas of two stocks overlap: the Western Baltic Spring Spawning Herring (WBSSH) and the Central Baltic Herring (CBH). Survey results of length‐at‐age data indicate in Subdivision (SD) 24, which is a part of the WBSSH management area, that a considerable fraction of CBH is present and correspondingly erroneously allocated to WBSSH stock indices. Accordingly, a stock Separation Function (SF) based on growth parameters was established to identify the fraction of CBH in the WBSSH area. In the present study, the SF was applied to 8 years of data from the GERman Acoustic Survey (GERAS), which is conducted annually in autumn in ICES subdivisions 21‐24. Results showed a distinct fraction of CBH in SD 24, and exclusion of the CBH greatly improved the quality of the GERAS index used in the assessment of the WBSSH stock.     

Laboratory studies on food intake, growth and food conversion of young herring, Clupea harengus (L.)

Food intake, growth and food conversion of young, O-group herring were studied at two temperatures and feeding regimes over a period of 19 weeks. The food intake of fish fed to satiation twice daily showed considerable variation. Food intake per fish at 14.5° C was about three times that at 6.5° C, and was generally much higher than in most other species of fish studied. The mean increase in wet weight over the 19-week period was 0.581 g/week at 14.5° C and 0.236 g/week at 6.5° C in fish fed to satiation and 0.094 g/week at 6.2° C and a ration of 1.3% of the body weight. Growth depensation was found to occur even in fish fed to satiation. The changes in specific growth rate, that is the percentage increase in weight/day, showed similar trends at different temperatures and food regimes. The mean conversion efficiency of fish on a ration of 1.3% at 6.2° C was higher than that of fish fed until satiation, at 14.5 and 6.5° C. The conversion efficiency of fish fed to satiation at 14.5° C showed a distinct decrease with increasing weight while at 6.5° C such a clear trend was not observed. In general, the conversion efficiency of young herring were found to be much lower than that of most other species studied. The weight exponent of the quantitative relationship between food intake and body weight at 14.5° C was 0.744. The total metabolic expenditure at 14.5° C, calculated using Winberg's (1956) 'utilization coefficient, gave a weight exponent of 0.773.     

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.     

Dispersion and mortality of yolk-sac herring (Clupea harengus L.) larvae from a spawning ground to the west of the Outer Hebrides

A patch of herring larvae located within 24 h of hatching froma spawning bed on the shelf west of the Outer Hebrides, wasfollowed for 7 days. Dispersion of the larvae was modelled usingsimple turbulent diffusion theory. The mortality rate, estimatedfrom the rate of decrease in the total population size, was–2% day^–1 over the tracking period which is verymuch less than rates calculated for herring larvae in otherareas by previous investigators. Successive observations ofthe length, weight, and the incidence of food in the intestinesof larvae, indicated that feeding commenced 5 days after hatchingand that the growth rate was 0.17 mm day^–1.     

Spatially structured interactions between a migratory pelagic predator, the Norwegian spring-spawning herring Clupea harengus L., and its zooplankton prey

Spring-spawning herring Clupea harengus was patchily distributed over large parts of the Norwegian Sea in May 1995–2005, during the early phase of the annual feeding migration. Overall, herring tended to be found in areas with intermediate biomasses of zooplankton prey, intermediate water temperatures and relatively high salinities. Herring had more food in their stomachs in areas of relatively low water temperature and high herring abundance. Hydrographical conditions revealed that herring was feeding mainly within Atlantic water masses, and more intensely in western and northern regions of the Norwegian Sea. Zooplankton biomass was patchily distributed, and was generally higher towards the western parts of the Norwegian Sea. Here, zooplankton biomass in year _{i +1} was also negatively associated with herring spawning stock biomass in year _i , while there was no evidence for such an association in the eastern region; indicating that herring may have a geographically structured 'top–down' effect on the recruitment of its zooplankton prey. The fact that herring was not typically associated with the areas containing the greatest zooplankton biomasses may reflect that the fish had not yet reached the most profitable feeding grounds or alternatively that herring was depleting zooplankton biomass.