Purification and characterization of pepsins from the arctic fish capelin (Mallotus villosus)

Two pepsins from the stomach of the arctic fish capelin (Mallotus villosus) have been isolated and characterized. The purification was achieved by ammonium sulphate precipitation, ion exchange chromatography and gel filtration. Both pepsins resemble mammalian pepsins regarding pepstatin sensitivity, amino acid composition, stability and specific activity. The major capelin pepsin has optimum activity at significantly higher pH than is common for mammalian pepsins, and the optimum pH is different with different substrates. Both pepsins have relatively high activity at low temperatures. The pepsins have mol. wt of about 25,000 which is significantly lower than that of mammalian pepsins.     

Incubation Temperature Effect on Qualitative and Quantitative Composition of Abnormalities and Mortality Rate in Embryogenesis of the Barents Sea Capelin Mallotus villosus (Osmeridae)

Journal of Ichthyology - The development of capelin Mallotus villosus from insemination to total resorption of the yolk is investigated at 5 and 8°C by the assessment of development of each...     

Genetic differentiation among populations of capelin Mallotus villosus from the West Coast of Greenland

Three populations of capelin, Mallotus villosus , from the west coast of Greenland were analyzed for variation among twelve proteins, determined by fifteen loci, using starch gel electrophoresis. The level of polymorphic loci at the 0.99 level varies from 0.39 to 0.62 and the heterozy gosity from 0.155 to 0.174. A significant heterogeneity among samples is demonstrated. The test values for homogeneity vary from 17.231 to 28.597 with four degrees of freedom.

Zusammenfassung

Genetische Unterschiede in Populationen der Lodde (Mallotus villosus) vor der Westküste Grönlands
In drei Populationen der Lodde, Mallotus villosus , von der Westküste Grönlands wurde die genetische Variation an zwölf Proteinen untersucht, die von fünfzehn Genorten bestimmt werden. Der Polymor-phiegrad (0.99 Kriterium) varüerte zwischen 0.39 und 0.62 und der Heterozygotiegrad zwischen 0.155 und 0.174. Bei den Populationen bestanden signifikante Unterschiede in den Genhäufigkeiten. Im Homogenitätstest mit vier Freiheitsgraden variierten die Testwerte zwischen 17.231 und 28.597.

Résumé

Différentiation génétique parmi des populations de capelans (Mallotus villosus) de la côte ouest du Greenland
L'électrophorèse sur le gel d'amidon permet d'étudier chez trois populations de capelans, Mallotus villosus , de la côte ouest du Greenland douze protéines déterminées par quinze loci. Le degré de polymorphisme au niveau 0.99 varie de 0.39 a 0.62 et le niveau d'hétérozygotie de 0.155 ä 0.174. L'hétérogénéité significative entre les populations a été trouvée.     

Effects of alternative prey on predation intensity from herring Clupea harengus and sandeel Ammodytes marinus on capelin Mallotus villosus larvae in the Barents Sea

Stomach contents from lesser sandeel Ammodytes marinus and herring Clupea harengus caught at one and three stations, respectively, were analysed to investigate predation intensity on capelin Mallotus villosus larvae. Most capelin larvae were found in the anterior sections of the stomachs close to the oesophagus, indicating that rapid digestion rates of larvae reduced the abundance in the posterior sections. The anterior sections of the stomachs had alternating layers of either copepods or capelin larvae and krill. This shows that the individual predators switched between feeding on either copepods or on krill and capelin larvae. A total of 549 capelin larvae was found in 440 fish stomachs. Capelin larvae were found in 20% of the sandeel stomachs, and 24, 34 and 62% of the stomachs from the three herring stations, respectively. Many of the predator stomachs contained more than five larvae, and up to 28 larvae were found in one herring stomach. The herring appeared to prey selectively on the largest capelin larvae, and the number of larvae per stomach was much higher in stomachs where krill had recently been eaten than where copepods were dominant. The predation intensity measured in this study is much higher than reported in earlier studies, and it is suggested that the predators were feeding using a searching image, and that the concentrations of alternative prey (copepods and krill) varied around a switching threshold.     

Mitochondrial DNA variation in Pacific capelin (Mallotus villosus catervarius) from the Sea of Okhotsk, inferred from PCR-RFLP analysis

Four population samples of Pacific capelin Mallotus villosus catervarius (Pennant, 1784) from geographically distant localities in the Sea of Okhotsk, Tauy Bay and the eastern coast of the Sakhalin Island, were examined using PCR-RFLP analysis of three mitochondrial DNA regions (A8/A6/COIII/ND3, ND3/ND4L/ND4, and ND5/ND6). The nucleotide divergence of mtDNA sequences among the samples, as well as the analysis of geographic heterogeneity of the haplotype frequencies and quantitative estimation of genetic differentiation performed by means of AMOVA, showed that the samples examined belonged to one panmictic population. Genealogic analysis of the mtDNA variation structure was carried out. It was demonstrated that the high level of haplotype diversity (0.9639 +/- 0.00015) along with the low level of nucleotide diversity (0.003818 +/- 0.0000003) pointed to the exponential rate of population growth of the capelin from the Sea of Okhotsk, which rather recently in its evolutionary history faced the bottleneck effect.     

Some aspects of the feeding and behavior of larvae of the Barents Sea capelin Mallotus villosus villosus (Salmoniformes, Osmeridae) in experimental conditions

The results of experimental investigations on the feeding and behavior of larvae of the Barents Sea capelin Mallotus villosus villosus during their transition to exogenous feeding are presented. Data concerning the passage duration of nauplii of Artemia salina along the intestine, feeding intensity, the portion of feeding larvae, swimming speed, sinking speed of non-feeding individuals, distances of responses and escape, as well as certain features of the passage of food through the intestines are given.     

The distribution and biology of the cestode Eubothrium parvum in capelin, Mallotus villosus, (Pallas) in the Barents Sea, and its use as a biological tag

Samples of Eubothrium parvum were obtained from capelin Mallotus villosus at 55 stations throughout the Barents Sea and from Balsfjord, North Norway. The parasite is distributed widely throughout the Barents Sea, but both incidence and intensity of infection are higher in the regions off Murmansk and the Kola peninsula, and Spitsbergen. E. parvum exhibits a seasonal peak in maturation and probably also in acquisition of new infections. The incidence of infection is greatest in 1 + fish, whereas the intensity is more independent of host age. It is suggested that the parasite requires only a single intermediate host, a plank-tonic copepod, and its distribution in relation to age of host is a reflection of the dietary preference shown by young capelin for copepods. The frequency distribution of E. parvum in capelin was over-dispersed in Balsfjord, where infection levels of between 1 and 28 parasites per fish were encountered in all samples, but under-dispersed in the Barents Sea, where infections of more than four parasites per fish were never found and even infections with three and four parasites were very local. It is suggested that the underdispersion is due to a very low probability of infection in the open waters of the sea. Although the presence of E. parvum cannot be used as a biological tag for capelin, its abundance and frequency distribution can. The difference in frequency distribution and the failure to find any heavily infected fish in the Barents Sea confirm the suggestion that the capelin of Balsfjord form a local isolated population, which does not migrate into the Barents Sea. The differences in infection levels within the Barents Sea suggest the further possibility that there are at least two stocks of capelin there, but this requires further investigation and confirmation.     

Invading predatory crustacean Dikerogammarus villosus eliminates both native and exotic species

As the tempo of biological invasions increases, explanations and predictions of their impacts become more crucial. Particularly with regard to biodiversity, we require elucidation of interspecific behavioural interactions among invaders and natives. In freshwaters in The Netherlands, we show that the invasive Ponto-Caspian crustacean amphipod Dikerogammarus villosus is rapidly eliminating Gammarus duebeni, a native European amphipod, and Gammarus tigrinus, until now a spectacularly successful invader from North America. In the laboratory, survival of single (unguarded) female G. duebeni was significantly lower when male D. villosus were free to roam as compared with isolated within microcosms. In addition, survival of paired (guarded) female G. duebeni was significantly lower when male D. villosus as compared with male G. duebeni were present. D. villosus killed and consumed both recently moulted and, unusually, intermoult victims. Survival of G. tigrinus was significantly lower when D. villosus were free to roam as compared with isolated within microcosms and, again, both moulted and intermoult victims were preyed upon. Male D. villosus were significantly more predatory than were females, while female G. tigrinus were significantly more often preyed upon than were males. Predation by D. villosus on both species occurred over a range of water conductivities, an environmental feature previously shown to promote amphipod coexistence. This predatory invader is predicted to reduce further the amphipod diversity in a range of freshwater habitats in Europe and North America.     

Mitochondrial DNA variation in Pacific capelin (<Emphasis Type="Italic">Mallotus villosus catervarius</Emphasis>) from the Sea of Okhotsk,inferred from PCR-RFLP analysis

Four population samples of Pacific capelin Mallotus villosus catervarius (Pennat, 1784) from geographically distant localities in the Sea of Okhotsk, Tauy Bay and the eastern coast of the Sakhalin Island, were examined using PCR-RFLP analysis of three mitochondrial DNA regions (A8/A6/COIII/ND3, ND3/ND4L/ND4, and ND5/ND6). The nucleotide divergence of mtDNA sequences among the samples, as well as the analysis of geographic heterogeneity of the haplotype frequencies and quantitative estimation of genetic differentiation performed by means of AMOVA, showed that the samples examined belonged to one panmictic population. Genealogic analysis of the mtDNA variation structure was carried out. It was demonstrated that the high level of haplotype diversity (0.9639 ± 0.00015) along with the low level of nucleotide diversity (0.003818 ± 0.0000003) pointed to the exponential rate of population growth of the capelin from the Sea of Okhotsk, which rather recently in its evolutionary history faced the bottleneck effect.     

Chemotaxonomic study on Thymus villosus from Portugal

The composition of the essential oils of four populations of Thymus villosus subsp. lusitanicus (Boiss.) Coutinho from Portugal was investigated by GC and GC-MS. To study the chemical polymorphism the results obtained from GC analyses of the volatile oils from individual plants from four populations were submited to Principal Component and Cluster analyses. A comparision with the essential oil of T. villosus subsp. villosus, previously studied by us was done. Important differences with regard to the major constituents in these two taxa were found. Linalool, geranyl acetate, geraniol and terpinen-4-ol were the main components of the essential oils of T. villosus subsp. lusitanicus, whereas in the oil of T. villosus subsp. villosus p-cymene, myrcene and alpha-terpineol were the major ones. Although, both taxa showed chemical polymorphism, different types of essential oils were characterized in each one: linalool; linalool/ terpinen-4-ol/trans-sabinene hydrate; linalool/1,8-cineole; geranyl acetate/geraniol; geranyl acetate/geraniol/1,8-cineole in T. villosus subsp. lusitanicus and p-cymene/camphor/linalool; p-cymene/borneol; linalool/geraniol/geranyl acetate; alpha-terpineol/camphor/myrcene in T. villosus subsp. villosus. Thus, the two subspecies of T. villosus can be easely differenciated by the composition of their essential oils.     

Comparison of liver enzymes in osmerid fishes: key differences between a glycerol accumulating species,rainbow smelt (Osmerus mordax), and a species that does not accumulate glycerol,capelin (Mallotus villosus)

Activities of enzymes associated with glycerol synthesis were compared in the liver of two osmerid fishes, the smelt (Osmerus mordax), which can accumulate high (400 mM) levels of glycerol and capelin (Mallotus villosus) that does not accumulate glycerol. Animals were sampled at approximately the same time of year and temperature thus negating potential seasonal effects. These species are closely related, reducing interpretative issues involving comparison between unrelated species. We found that key enzyme activities were elevated in the smelt relative to the non-glycerol accumulating capelin, namely enzymes involved with glycolysis (phosphofructose kinase-1 and aldolase), amino acid metabolism (aspartate aminotransferase and alanine aminotransferase), gluconeogenesis (phosphoenolpyruvate carboxykinase) and glycerol synthesis (glycerol-3-phosphate dehydrogenase). The enzyme profiles strongly support the hypothesis that smelt can synthesize glycerol by utilizing glycogen and amino acids as the carbon source and that they have increased capacity for metabolic flux through loci required for synthesis of the three carbon intermediate dihydroxyacetone phosphate and subsequently glycerol synthesis.     

Shrinkage correction and length conversion equations for Theragra chalcogramma, Mallotus villosus and Thaleichthys pacificus

Preservation of walleye pollock Theragra chalcogramma, capelin Mallotus villosus and eulachon Thaleichthys pacificus by freezing decreased fork length ( L _F) up to 1·8, 5·6 and 2·7% and reduced mass by up to 8·4, 3·5 and 1·1%, respectively. Shrinkage of walleye pollock standard length ( L _S) was greater for fish in 95% ethanol v . 5% formalin and for fish in 10% formalin v . frozen. Equations describing the shrinkage and loss in mass for these species are presented as well as conversions between different length measurements ( L _S, L _F and total length, L _T) for fishes that were frozen.     

Identifying the spatial scale of density-dependent interaction of predators with schooling fish in the southern Labrador Current

Previous work along the coast of Newfoundland has shown that aggregative responses of puffins, Fraiercula arctica , and common guillemots, Uria aalge , to schooling fish, primarily capelin, Mallotus villosus , occur at spatial scales of 2-6 km. The present work tests the hypothesis that coastal upwelling results in aggregation of capelin at this scale, and that the scale of interaction with predators can be predicted from wind-driven coastal upwelling. An observational experiment in 1987 showed that a relatively simple physical model could be used to predict the spatial scale of upwelling, that schooling fish aggregate at this scale after upwelling, that aggregation of schooling fish at the scale of upwelling decays as upwelling relaxes, and that the spatial association of puffins and guillemots with schooling fish reaches a maximum at this scale. These results suggest that relatively simple physical theory can, in some cases, be used to identify the spatial scale of interaction of predators with prey.     

The ability of gadoids to take advantage of a short-term high availability of forage fish: the example of spawning aggregations in Barents Sea capelin

During 11 March to 4 April 2002, the distribution of Barents Sea capelin Mallotus villosus along the coast of Finnmark, northern Norway, was covered four times by combining acoustic survey with trawling, synoptically and simultaneously sampling capelin and its main fish predators; cod Gadus morhua, haddock Melanogrammus aeglefinus and saithe Pollachius virens. The surveys demonstrated how these gadoid predators were able to exploit such a short‐term abundance of forage fish. The predator aggregation as well as the stomach fullness and proportion of capelin in their diet followed the capelin spawning migration, increasing in areas and periods with increasing capelin abundance. Capelin clearly constituted most of the biomass in stomachs of cod (97%), haddock (87%) and saithe (96%). The stomach fullness was highest in cod and lowest in haddock, although in areas with low capelin abundance, saithe had more capelin in their stomachs. The total length (L_T) of capelin in predator stomachs increased with predator L_T, but the proportion of capelin in the diet was not influenced by predator L_T. The capelin in predator stomachs was significantly smaller than capelin in the trawl hauls, also when compared within the same sex, indicating feeding selectivity towards weaker individuals. Female capelin, being significantly smaller than the males, predominated in the diet of haddock, whereas in cod and saithe the sex ratio was more equal. Male capelin predominated in the predator diet during the pre‐spawning period, whereas the females predominated as the spawning commenced. During the overall study period, most of the female capelin in predator stomachs was in a pre‐spawning or a spawning stage, whereas the majority of the males appeared to be spent. Regardless of sex, the percentage of spent, relative to pre‐spawning or spawning capelin in the diet of the predators, followed the capelin spawning dynamics, increasing with time as the spawning progressed.     

Egg cannibalism in Barents Sea capelin in relation to a narrow spawning distribution

The spawning distribution of Barents Sea capelin Mallotus villosus off northern Norway in 2002 was restricted to a narrow area at the easternmost spawning grounds. There was an increase in abundance and a marked shift in aggregation area from east towards west during 11 March to 4 April, as the capelin went from prespawning through spawning to spent stages. The capelin stomachs were either empty or contained capelin eggs, often mixed with sand. Both the occurrence of cannibals and stomach fullness increased with stage of maturity, being highest in spent fish, and higher in males than in females. No differences were found between cannibals and non-cannibals of the same sex when adjusted for total length ( L _T), age, L _T at age and condition factor. At similar stomach fullness, the females had consumed more eggs than the males. This was attributed to a higher proportion of empty and broken eggs and less sand in females compared to males. In the full stomachs, the mean consumption in females and males was 623 and 334 eggs, respectively, whereas the respective maximum egg consumption was 871 and 683 eggs. The minimum estimates (given no digestion) of mean ± 95% CL egg consumption in spent females and males were 75·4 ± 6·9 and 58·4 ± 12·0 eggs, respectively, implying a minimum mortality of 1–2% of the total egg production caused by cannibalism.     

Atlantic capelin (Mallotus villosus) tetranucleotide microsatellites

Twelve microsatellite loci developed for Atlantic capelin (Mallotus villosus) using magnetic bead hybridization enrichment for tetranucleotide microsatellites revealed five loci composed of single repeat elements and six composed of complex repeats. Forty‐four beach‐spawning females from three different northwestern Atlantic Newfoundland beach‐spawning populations were screened at each locus. Loci were polymorphic (two to 59 alleles per locus) and all but two exhibited high heterozygosity (0.86–1). The loci are considered suitable for addressing questions related to fine‐scale population structure, spawning fidelity and survivorship/kinship issues.     

The foraging decisions of a central place foraging seabird in response to fluctuations in local prey conditions

During reproduction, seabirds need to balance the demands of self- and offspring-provisioning within the constraints imposed by central place foraging. To assess behavioral adjustments and tolerances to these constraints, we studied the feeding tactics and reproductive success of common murres (also known as common guillemots) Uria aalge , at their largest and most offshore colony (Funk Island) where parents travel long distances to deliver a single capelin Mallotus villosus to their chicks. We assessed changes in the distance murres traveled from the colony, their proximate foraging locations and prey size choice during two successive years in which capelin exhibited an order of magnitude decrease in density and a shift from aggregated (2004) to dispersed (2005) distributions. When capelin availability was low (2005), parental murres increased their maximum foraging distances by 35% (60 to 81 km) and delivered significantly larger capelin to chicks, as predicted by central place foraging theory. Murres preferred large (>140 mm) relative to small capelin (100–140 mm) in both years, but unexpectedly this preference increased as the relative density of large capelin decreased. We conclude that single prey-loading murres target larger capelin during long foraging trips as parents are 'forced' to select the best prey for their offspring. Low fledgling masses suggest also that increased foraging time when capelin is scarce may come at a cost to the chicks (i.e. fewer meals per day). Murres at this colony may be functioning near physiological limits above which further or sustained adjustments in foraging effort could compromise the life-time reproductive success of this long-lived seabird.     

Growth of the Barents Sea capelin,Mallotus villosus,in relation to climate

Synopsis In order to investigate a possible relationship between temperature and fish length growth in the Barents Sea capelin stock, estimates of environmental temperature in the feeding season were compared to estimates of length growth during the same season. The mean temperature of the capelin feeding area was calculated by averaging the temperatures in September for the depth interval 10–200 m in statistical rectangles. The estimates of capelin growth were obtained for the same rectangles using backcalculation of length from otoliths. Correlation coefficients for the relationship between water temperature and growth were 0.70 and 0.53 for two- and three-year-olds, respectively, when all the material was considered, and between 0.85 and 0.91 for within-year data. In addition, a close correspondence between feeding area and growth rates was found.     

The timing of spawning in capelin (Mallotus villosus Müller) at a coastal location in eastern Newfoundland

We tested three hypotheses concerning the timing of spawning for a circumpolar species, capelin (Mallotus villosus), for which timing of larval emergence is known to be synchronized by physical conditions. The first hypothesis, developed from previous studies, was that spawning would be synchronized by upwelling events. Initial results from Middle Cove Beach in eastern Newfoundland indicated that spawning was not synchronized with upwelling. We next hypothesized that spawning was a function of several environmental variables. Results from logistic regression indicated that neither single-factor nor multi-factor models could explain the timing of spawning. Single variables could predict spawning in some years but no variable could reliably predict the time of spawning year after year. Finally, we hypothesized that the probability of spawning increased as a set of significant variables approached preferred levels. For capelin at Middle Cove, the set of variables that influence capelin spawning were identified as wave height, sea surface roughness and capelin abundance in the water. Thus only a combination of variables explained the timing of spawning for capelin. Preferred conditions for capelin spawning were wave heights less than 20 cm at the beach, a sea surface with a slight ripple, and an intermediate rank abundance of capelin in the water corresponding to hundreds to thousands of individuals. Capelin abundance alone was not a useful predictor. During the course of the study we observed a shift in the dates that capelin arrived and spawned at the beach. During 1987–1990 capelin spawned at Middle Cove Beach during June, but in more recent years (1991–1993) capelin did not arrive or spawn until July.