Nucleotide sequence variation of the mitochondrial COI gene in several eelpout species of the genus Zoarces (Zoarcidae, Pisces)

The mitochondrial COI gene was for the first time sequenced in eelpout species of the genus Zoarces from the Taui Bay of the Sea of Okhotsk: notched-fin Z. elongatus Kner, 1868 and the new sympatric form Zoarces sp., differing in a large set of morphological traits. The two species were compared with European eelpout Z. viviparus Linnaeus, 1758 from the Finnish Bay of the Baltic Sea. Divergence and phylogenetic analyses demonstrated a greater genetic similarity between Zoarces sp. and Z. viviparous than between sympatric Zoarces sp. and Z. elongatus.     

Restriction fragment variation of the mitochondrial gene of cytochrome b in some taxa of the Eelpout family (pisces, perciformes, zoarcidae) from the northern Sea of Okhotsk

The first analysis of restriction fragment length polymorphism (RFLP) of the mitochondrial gene of cytochrome b of fish from the Eelpout Family (Zoarcidae) from the northern Sea of Okhotsk has been performed. The mean genetic distances between the subfamilies Zoarcinae and Lycodinae, subfamilies Lycodinae and Gymnelinae, and subfamilies Zoarcinae and Gymnelinae are 11.14, 13.87, and 14%, respectively. Species of the genus Lycodes (Lycodinae) are characterized by a mtDNA small divergence (on average, 2.04%) and are divided into two groups on the basis of lateral line morphology. Magadania skopetsi, an endemic of Tauiskaya Bay of the Sea of Okhotsk, is genetically close to Hadropareia middendorffii from the same subfamily (Gymnelinae). The mtDNA sequences of species from the genus Lycogrammoides (Lycodinae) differ by 3.83%. The genetic differences between the eelpout Zoarces elongatus Zoarces sp. (d = 4.54%) suggest that the latter is a separate species.     

Relationships of eelpouts of genus Zoarces (Zoarcidae, Pisces) inferred from molecular genetic and morphological data

Molecular genetic and morphological analysis of eelpouts of the genus Zoarces was carried out. Based on the mitochondrial DNA sequence variation, haplotypes of notched-fin eelpout, Z. elogatus, more closely related Fedorov eelpout, Z. fedorovi, and common eelpout, Z. viviparus, as well as of Andriyashev eelpout, Z. andriashevi, were grouped in one macrocluster. Haplotypes of American eelpout, Z. americanus, and blotched eelpout, Z. gillii, clustered separately from other species. The genetic differences between Z. gillii and the other eelpout species were very high for within-genus comparisons, constituting 7.62%. Species divergence in terms of morphological characters was generally consistent with molecular genetic data and confirmed distinct isolation of American eelpout, and especially of blotched eelpout.     

Restriction fragment variation of the mitochondrial gene of cytochrome b in some taxa of the Eelpout Family (Pisces, Perciformes, Zoarcidae) from the northern Sea of Okhotsk

The first analysis of restriction fragment length polymorphism (RFLP) of the mitochondrial gene of cytochrome b of fish from the Eelpout Family (Zoarcidae) from the northern Sea of Okhotsk has been performed. The mean genetic distances between the subfamilies Zoarcinae and Lycodinae, subfamilies Lycodinae and Gymnelinae, and subfamilies Zoarcinae and Gymnelinae are 11.14, 13.87, and 14.00%, respectively. Species of the genus Lycodes (Lycodinae) are characterized by a mtDNA small divergence (on average, 2.04%) and are divided into two groups on the basis of lateral line morphology. Magadania skopetsi, an endemic of Tauiskaya Bay of the Sea of Okhotsk, is genetically close to Hadropareia middendorffii from the same subfamily (Gymnelinae). The mtDNA sequences of species from the genus Lycogrammoides (Lycodinae) differ by 3.83%. The genetic differences between the eelpout Zoarces elongatus Zoarces sp. (d = 4.54%) suggest that the latter is a separate species.     

Thermal physiology of the common eelpout (Zoarces viviparus)

We investigated the temperature dependence of some physiological parameters of common eelpout (Zoarces viviparus) from different locations (North Sea, Baltic Sea and Norwegian Sea) on acclimation temperature (3 degrees C and 12 degrees C) and acute temperature variation. The lethal limit of 12 degrees C-acclimated eelpout was determined as the critical thermal maximum [loss of equilibrium (LE) and onset of muscular spasms (OS)] and it was found to be 26.6 degrees C for LE and 28.8 degrees C for OS for all populations. However, these parameters do not have any relevant ecological interpretation. We therefore investigated the effect of gradually increased water temperature on standard metabolic rate (measured as resting oxygen consumption Mo2) and critical oxygen concentration ([O2]c) of eelpouts. Acclimation to low temperature (3 degrees C) resulted in partial compensation of Mo2, paralleled by a decrease of activation energy for Mo2 (from 82 kJ mol(-1) at 12 degrees C to about 50 kJ mol(-1) at 3 degrees C) in North Sea and Baltic Sea eelpouts. At the same time, Norwegian eelpout showed no acclimation of oxygen demand to warm temperature (12 degrees C) at all. The scope for eelpout aerobic metabolism shrank considerably with increased acclimation temperature, as [O2]c approached water oxygen concentrations. At 22.5+/-1 degrees C the [O2]c reached air saturation, which is equivalent to the upper critical temperature (TcII) and at this temperature the aerobic scope for the metabolism completely disappeared. In line with previous insight, the comparative analysis of the temperature dependence of Mo2 of Z. viviparus from different populations suggests that a pejus (sub-critical) temperature for this species is about 13-15 degrees C. In conclusion, the capacity to adjust aerobic metabolism relates to thermal tolerance and the bio-geographical distribution of the species. Global warming would thus be likely to cause a shift in the distribution of this species to the North.     

Panmixia in Zoarces viviparus: implications for environmental monitoring studies

In this study, the genetic population structure of the eelpout Zoarces viviparus was investigated by using microsatellites. Samples were collected at 10 sites in the Baltic Sea, covering a distance of c. 90 km. Ten newly developed microsatellite loci were used to infer the population structure. No global spatial genetic differentiation was found (global F(ST) = 0·0001; D(est) = -0·0003), indicating strong gene flow at this scale, nor any clear pattern of isolation by distance. The results suggest that gene flow among the studied populations of Z. viviparus is stronger than usually thought, which might be caused by environmental homogeneity. This is important for planning and evaluating monitoring activities in this species and for the interpretation of ecotoxicological studies. Strong migration might lead to wrong conclusions concerning the pollution in a given area. Therefore, reference stations should be placed at a larger distance than presently practiced.     

Effects of seasonal and latitudinal cold on oxidative stress parameters and activation of hypoxia inducible factor (HIF-1) in zoarcid fish

Acute, short term cooling of North Sea eelpout Zoarces viviparus is associated with a reduction of tissue redox state and activation of hypoxia inducible factor (HIF-1) in the liver. The present study explores the response of HIF-1 to seasonal cold in Zoarces viviparus, and to latitudinal cold by comparing the eurythermal North Sea fish to stenothermal Antarctic eelpout (Pachycara brachycephalum). Hypoxic signalling (HIF-1 DNA binding activity) was studied in liver of summer and winter North Sea eelpout as well as of Antarctic eelpout at habitat temperature of 0°C and after long-term warming to 5°C. Biochemical parameters like tissue iron content, glutathione redox ratio, and oxidative stress indicators were analyzed to see whether the cellular redox state or reactive oxygen species formation and HIF activation in the fish correlate. HIF-1 DNA binding activity was significantly higher at cold temperature, both in the interspecific comparison, polar vs. temperate species, and when comparing winter and summer North Sea eelpout. Compared at the low acclimation temperatures (0°C for the polar and 6°C for the temperate eelpout) the polar fish showed lower levels of lipid peroxidation although the liver microsomal fraction turned out to be more susceptible to lipid radical formation. The level of radical scavenger, glutathione, was twofold higher in polar than in North Sea eelpout and also oxidised to over 50%. Under both conditions of cold exposure, latitudinal cold in the Antarctic and seasonal cold in the North Sea eelpout, the glutathione redox ratio was more oxidised when compared to the warmer condition. However, oxidative damage parameters (protein carbonyls and thiobarbituric acid reactive substances (TBARS) were elevated only during seasonal cold exposure in Z. viviparus. Obviously, Antarctic eelpout are keeping oxidative defence mechanisms high enough to avoid accumulation of oxidative damage products at low habitat temperature. The paper discusses how HIF could be instrumental in cold adaptation in fish.     

Variations in antifreeze activity and serum inorganic ions in the eelpout Zoarces viviparus: antifreeze activity in the embryonic state.

The eelpout Zoarces viviparus is a common inhabitant in the shallow waters along the Danish coastline. Specimens were caught in the brackish (12-16 per thousand) Roskilde fjord where water temperatures range from >20 degrees C during summer to subzero in winter. The serum melting points found in Z. viviparus varied between -0.76 (September) to -0.94 degrees C (January). Eighty to 97% of the serum melting points could be attributed to sodium, chloride and potassium. Hysteresis freezing points showed seasonal variation varying from -0.83 (September) to -2.08 degrees C (February). Serum antifreeze activity showed a seasonal variation with high levels (>1.2 degrees C) in winter and low levels (<0.1 degrees C) during summer and autumn. Antifreeze proteins are responsible for this antifreeze activity. Antifreeze activity was also found in Z. viviparus during their embryological development in the female ovary. Embryo thermal hysteresis reached the maximum level (approx. 0.6 degrees C) during December and maintained this level until parturition in January. Antifreeze activity seems unaffected by diminishing ice crystal fractions at ice fractions below 0.1 whereas ice fractions above 0.1 caused a decline in antifreeze activity.     

Features of Transition from Larva to Juvenile in Fishes with Different Types of Early Ontogeny

The transition from the larva state to the juvenile state (i.e. morphological condition characterised by mainly adult characters) was examined in three marine fish species: herring Clupea pallasi marisalbi, wolffish Anarhichas lupus, and eelpout Zoarces viviparus, based on external morphology and skeletal development. In spite of the different reproductive styles (oviparity, facultative viviparity, and obligate viviparity, respectively) and different types of early ontogeny (indirect, transitory, and direct, respectively), the beginning of the juvenile state occurred at similar total lengths (TL), which were approximated as 35 mm TL in herring and eelpout, and 32 mm TL in wolffish. Features of ontogeny were compared, assuming that the beginning of the juvenile state represented an uniform characteristic of morphological development for these species. It was proposed that the beginning of the larva or juvenile periods (sensu Balon) could not coincide with the beginning of the larva and juvenile states in the ontogeny of some species.     

Trends in abundance of benthic and demersal fish populations of the lower Forth Estuary, East Scotland, from 1982–2001

The benthic and demersal fishes of the lower Forth Estuary, east Scotland, were sampled by Agassiz trawling from January 1982–December 2001. Three stations of varying distance from the mouth of the estuary were trawled at high and low water in 5 months of each year, a total of 30 trawls per annum using the same ship and net design throughout. Of 30 species collected over this period, whiting Merlangius merlangus was the most abundant, followed by eelpout Zoarces viviparus , pogge Agonus cataphractus , flounder Platichthys flesus and plaice Pleuronectes platessa . The 10 most abundant species accounted for 97·8% of the total benthic-demersal fish abundance. There was no trend in total annual species richness sampled, though total annual abundance of fishes captured decreased significantly from 1982–2001. This was largely caused by significant declines in abundance of whiting and eelpout. Fatherlasher Myoxocephalus scorpius showed a significant linear increase. None of the seven remaining most abundant species exhibited significant trends in abundance. Decline of eelpout may be attributable to increases in water temperature adversely affecting reproductive success. This is suggested by a significant negative correlation of mean annual abundance with January bottom temperature in the previous year.     

Comparative osteology,relationships, and systematics of fish of the genus <Emphasis Type="Italic">Zoarces</Emphasis> (Zoarcidae,Perciformes)

Comparative-osteological investigation of all species of the genus Zoarces is made: Z. fedorovi, Z. andriashevi, Z. elongatus, Z. viviparus, Z. americanus, and Z. gillii. In the skeleton of hyoid arch in species of Zoarces, Lycozoarces, and Lycodes, paired bony elements are found, not described previously, identified as parurohyalia which may be synapomorphy of Zoarcidae. Differences between the Zoarces species are revealed by 56 external morphological and osteological characters. Cladistic analysis yielded one tree 135, Ci 0.57, Ri 0.54 long. Cladistic analysis confirms validity of a recently described species Z. fedorovi, which turned out to be the most generalized in the genus Zoarces, as well as the validity of Z. elongatus. The species Z. americanus and Z. gillii occupy a terminal position preventing their isolation to independent genera as was previously supposed, as this would lead to paraphyly of the genus Zoarces.     

Features of the biology of the eelpout <Emphasis Type="Italic">Zoarces fedorovi</Emphasis> (Perciformes: Zoarcidae) from Tauisk bay,Sea of Okhotsk

The results of long-term research on the biology of the viviparous Fedorov eelpout Zoarces fedorovi from the Tauiskaya Guba in the Sea of Okhotsk are generalized. It was determined that in winter and spring this species inhabits strongly freshened sea areas and in summer and fall it inhabits, as a rule, estuarine areas in the tide effect zone. Z. fedorovi did not leave the coastal zone, even during the ebb, and could remain in colder fresh water up to 12 h a day. Data on the age, sexual and size structure of the catches, character of linear growth, increase in body mass and fecundity, size, as well as the growth rates of the larvae and fry during prenatal development of Z. fedorovi are cited.     

Aerobic mitochondrial capacities in Antarctic and temperate eelpout (Zoarcidae) subjected to warm versus cold acclimation

Capacities and effects of cold or warm acclimation were investigated in two zoarcid species from the North Sea (Zoarces viviparus) and the Antarctic (Pachycara brachycephalum) by investigating temperature dependent mitochondrial respiration and activities of citrate synthase (CS) and NADP⁺ -dependent isocitrate dehydrogenase (IDH) in the liver. Antarctic eelpout were acclimated to 5°C and 0°C (controls) for at least 10 months, whereas boreal eelpout, Z. viviparus (North Sea) were acclimated to 5°C and to 10°C (controls). Liver sizes were found to be increased in both species in the cold, with a concomitant rise in liver mitochondrial protein content. As a result, total liver state III rates were elevated in both cold-versus and warm-exposed P. brachycephalum and Z. viviparus, with the highest rates in boreal eelpout acclimated to 5°C. CS and IDH activities in the total liver were similar in Z. viviparus acclimated to 5°C and 10°C, but decreased in those warm acclimated versus control P. brachycephalum. Enzyme capacities in the total liver were higher in eelpout from Antarctica than those from the North Sea. In conclusion, cold compensation of aerobic capacities in the liver seems to be linked to an increase in organ size with unchanged specific mitochondrial protein content. Despite its life in permanently cold climate, P. brachycephalum was able to reduce liver aerobic capacities in warm climate and thus, displayed a capacity for temperature acclimation.     

Spatial and length distribution,sex composition,and feeding of the notched-fin eelpout <Emphasis Type="Italic">Zoarces elongatus</Emphasis> (Perciformes: Zoarcidae) in the Sea of Okhotsk

Based on the data from six bottom trawl surveys carried out on the shelf of the Sea of Okhotsk in 1997–2007, the features of the distribution, size and sex structure, and food composition of the notched-fin (east) eelpout Zoarces elongatus were analyzed. It was established that this species regularly occurs in trawl catches only in the eastern part of the sea and that the density of its schools decreases with depth. It was recorded that females of the notched-fin eelpout are larger than males. The sex ratio of the fishes varies from year to year. Females prevail at depths less than 25 m and males prevail at depths greater than 25 m.     

Mortality of fish from the by-catch of shrimp vessels in the North Sea

The results from 3 different rogrammes concerning by-catch mortality in shrimp fisheries are reported. Mortalities increased consigrably after the catch passed the sorting sieve. 100 % mortality was detected for whiting ( Merlangius merlangus ). Mortalities were about 10% for sculpin ( Myxocephalus scorpius ), hooknose ( Agonus catapbractus ), and eelpout ( Zoarces viviparus ) in the discard fraction. Survival of flatfish depended strongly on the species, the size of the secimens as well as the catch and catch processing conditions, and ranged from 17 to 100%. No difirences could be detected in the survival after sorting on different machines. However, due to better sorting efficiency, the rotary sieve may reduce mortality of fish in the by-catch.

Zusammenfassung

Sterblichkeit von Fischen aus dem Beifang von Garnelenkuttern in tier Nordsee
Es werden die Ergebnisse von 3 verschiedenen Untersuchungsprogramrnen zur Beifansterblichkeit in der Garnelenfischerei mitgeteilt. Die Sterblichkeit nahm beträchtlich zu, wenn der Beikng das Sortiersieb passiert hatte. 100% Sterblichkeit wurde für Wittlinge ( Merlangius merlangus ) ermittelt. Für Seeskorpione ( Myoxocephalus scorpius ), Steinpicker ( Agonus cataphractus ) und Aalmuttern ( Zoarces viviparus ) lag die Sterblichkeit in der Discard-Fraktion iei 10%. Die Überlebensraten bei Plattfischen waren stark von der jeweilien Art, der Größe derTiere, wie auch vom Fang und der Fangbearbeitung abhängig und bewegten sici zwischen 17 und 100%. Es konnten keine Unterschiede in den Überlebensraten nach der Sortierung mit verschiedenen Maschinen festgestellt werden. Allerdins kann der Trommelsonierer aufgrund seiner besseren Sortiereigenschaften die Sterblichkeit im Beifang herabsetzen.     

Biology of eelpout <Emphasis Type="Italic">Zoarces elongatus</Emphasis> (Zoarcidae) from Tauysk Bay,the Sea of Okhotsk

Based on the material collected in Tauysk Bay, the Sea of Okhotsk, from 1997 to 2014, the features of spatial distribution, body length and body weight composition, sex ratio, and some aspects of growth rate and reproduction of eelpout Zoarces elongatus are described. The representatives of the species prefer the biotopes with the littoral composed of large stones or pebbles, mainly from the upper border of the intertidal zone to the depths between 10 and 20 m. The catches mainly include the individuals with the body length 300?400 mm and body weight 0.1?100.0 g at the age between 5+ and 6+. Maximum registered body length, body weight, and age are as follows: 560.0 mm, 742 g, and 11+. The sex ratio (males: females) is 1: 1.2. In Tauysk bay, the release of juveniles occurs from November to December, and fecundity ranges from 20 to 185 eggs.     

Strong isolation-by-distance in the absence of genetic population structure in the eelpout (Zoarces viviparus,Linnaeus 1758)

The eelpout (Zoarces viviparus) is a benthic marine fish, which has long been assumed to be philopatric. This made it particularly suitable for environmental monitoring programs as it allows matching its content of pollutants to the area of catch. However, a recent small scale genetic study in the Baltic Proper suggested that gene-flow might be stronger than generally believed and may occur frequently up to a distance of at least 90 km. As challenging the assumption of philopatry might have strong implications for environmental monitoring studies, we tested the hypothesis of philopatry at a larger geographical scale using ten microsatellite loci. A total of 220 individuals sampled from eight locations covering almost the entire geographic range of the species was genotyped. Our results show that genetic diversity decreases from the North Sea to the northeastern Baltic Sea. No strong population structuring was found, but a highly significant isolation-by-distance pattern was detected, suggesting a stepwise migration pattern among populations. Thus, the hypothesis of long-distance migration can be falsified. It is more likely that only limited gene flow exists among adjacent populations without any barriers between them. Our results suggest that dispersal in the eelpout is weak enough to retain this species as an important bioindicator. However, we suggest that reference stations should be placed in an appropriate distance to avoid misleading results from migrating individuals. We conclude that a more precise knowledge on migration rates of the eelpout is required in order to get more reliable insights into the potential area over which the concentration of environmental pollutants is integrated.