Complete nucleotide sequence and variation of mitochondrial DNA from 10 individuals of walleye pollock, Theragra chalcogramma

ABSTRACT:   The complete mitochondrial genome sequence of 10 walleye pollocks, Theragra chalcogramma , from the Japan Sea and Bering Sea was determined. The 16 568–16 571 bp genome contains the same 37 mitochondrial structural genes (two ribosomal RNA, 22 transfer RNA, and 13 protein-coding genes) as found in all other vertebrates analyzed, in an organization identical to that of other bony fish. The major non-coding region had several conserved sequence features. Nucleotide variations of ND1, ND5, and control region were high, and these regions appear to be good candidates for high-resolution markers in population studies.     

Spatial and temporal patterns of walleye pollock (Theragra chalcogramma) spawning in the eastern Bering Sea inferred from egg and larval distributions

Walleye pollock Theragra chalcogramma (pollock hereafter) is a key ecological and economic species in the eastern Bering Sea, yet detailed synthesis of the spatial and temporal patterns of pollock ichthyoplankton in this important region is lacking. This knowledge gap is particularly severe considering that egg and larval distribution are essential to reconstructing spawning locations and early life stages drift pathways. We used 19 yr of ichthyoplankton collections to determine the spatial and temporal patterns of egg and larval distribution. Generalized additive models (GAMs) identified two primary temporal pulses of pollock eggs, the first occurring from 20 February to 31 March and the second from 20 April to 20 May; larvae showed similar, but slightly lagged, pulses. Based on generalized cross‐validation and information theory, a GAM model that allowed for different seasonal patterns in egg density within three unique areas outperformed a GAM that assumed a single fixed seasonal pattern across the entire eastern Bering Sea. This ‘area‐dependent’ GAM predicted the highest densities of eggs (i.e., potential spawning locations) in three major areas of the eastern Bering Sea: near Bogoslof Island (February–April), north of Unimak Island and the Alaska Peninsula (March–April), and around the Pribilof Islands (April–August). Unique temporal patterns of egg density were observed for each area, suggesting that pollock spawning may be more spatially and temporally complex than previously assumed. Moreover, this work provides a valuable baseline of pollock spawning to which future changes, such as those resulting from climate variability, may be compared.     

Temperature-dependent stock-recruitment model for walleye pollock (Theragra chalcogramma) around northern Japan

Changes in fish year‐class strength have been attributed to year‐to‐year variability in environmental conditions and spawning stock biomass (SSB). In particular, sea temperature has been shown to be linked to fish recruitment. In the present study, I examined the relationship between sea surface temperature (SST), SSB and recruitment for two stocks of walleye pollock (Theragra chalcogramma) around northern Japan [Japanese Pacific stock (JPS) and northern Japan Sea stock (JSS)] using a temperature‐dependent stock‐recruitment model (TDSRM). The recruitment fluctuation of JPS was successfully reproduced by the TDSRM with February and April SSTs, and February SST was a better environmental predictor than April SST. In addition, the JPS recruitment was positively related to February SST and negatively to April SST. The JSS recruitment modeled by the TDSRM incorporating February SST was also consistent with the observation, whereas the relationship between recruitment and February SST was negative, that is the opposite trend to JPS. These findings suggest that SST in February is important as a predictor of recruitment for both stocks, and that higher and lower SSTs in February act favorably on the recruitment of JPS and JSS respectively. Furthermore, Ricker‐type TDSRM was not selected for either of the stocks, suggesting that the strong density‐dependent effect as in the Ricker model does not exist for JPS and JSS. I formulate hypotheses to explain the links between SST and recruitment, and note that these relationships should be considered in any future attempts to understand the recruitment dynamics of JPS and JSS.     

Geographical variations in infection by larval Anisakis simplex and Contracaecum osculatum (Nematoda, Anisakidae) in walleye pollock Theragra chalcogramma stocks off Hokkaido, Japan

ABSTRACT: Stocks of walleye pollock Theragra chalcogramma collected from: (i) the Sea of Japan (off Rebun Island and Kumaishi); (ii) the Pacific coast (off Shikabe and eastern Hokkaido); and (iii) Nemuro Strait off Hokkaido, northern Japan, were examined for anisakid nematodes during December 1999 to February 2000, and the prevalence and abundance of Anisakis simplex and Contracaecum osculatum larvae were compared among the various sampling sites for fish of the same size and age. Anisakis simplex was generally more abundant than C. osculatum . Infection by A. simplex varied between the aforementioned stocks of walleye pollock as well as within stocks, whereby fish from off Rebun Island and Nemuro Strait were infected the most, followed by those from off the Pacific coast and Kumaishi. Infection by C. osculatum differed between the host stocks, and C. osculatum was the most abundant among the fish from Nemuro Strait. The infection variations seemed to be due to differences in host growth rate, host feeding habit, and the distribution of marine mammal final hosts. The results indicate that these two larval nematodes are useful biological indicators for the population study of walleye pollock in Japanese waters.     

Interannual variability in hatching period and early growth of juvenile walleye pollock, Theragra chalcogramma, in the Pacific coastal area of Hokkaido

Juvenile walleye pollock of the Japanese Pacific population were collected from the Funka Bay [spawning ground; 16–64 mm fork length (FL)] in spring and the Doto area (nursery ground; 70–146 mm FL) in summer. Hatch dates were estimated by subtracting the number of otolith daily increments from sampling dates, and their early growth was back‐calculated using otolith radius–somatic length relationships. Interannual change of the hatching period was observed during 2000–02, and the peaks ranged from mid‐February in 2000 to early‐April in 2002. In 2000, when a strong year class occurred, early life history of the surviving juveniles could be characterized by early hatching and slower growth in the larval stage (<22 mm length). Higher growth rate in 2001 and 2002 did not always lead to good survival and recruitment success. Even though their growth was slow in 2000, the larvae hatched early in the season had larger body size on a given date than faster‐growing larvae hatched in later season in 2001 and 2002. Bigger individuals at a certain moment may have advantage for survival. The delay of hatching period may result in higher size‐selective mortality, and as a necessary consequence, back‐calculated growth in 2001 and 2002 could shift towards higher growth rate, although abundance of such a year class would be at the lower level. Variability in spawning period, early growth and their interaction might have a strong relation to larval survival through cumulative predation pressure or ontogenetic changes in food availability.     

Abiotic and biotic factors affecting recruitment variability of walleye pollock (Theragra chalcogramma) off the Pacific coast of Hokkaido,Japan

Abiotic and biotic factors affecting the recruitment variability of the Japanese Pacific stock (JPS) of walleye pollock (Theragra chalcogramma) were examined using a bivariate regression and multivariate combined model. Of the abiotic variables around Funka Bay (spawning ground), February sea surface temperature (SST) and wind direction index showed significant bivariate relationships with recruitment. February SST was positively related to recruitment, suggesting that warmer water temperature in February favors JPS recruitment. On the other hand, the relationship between February wind direction index and recruitment predicts high JPS recruitment under predominant northwest winds in February. For the biotic variables in the Doto area (nursery ground), significant and negative bivariate relationships with recruitment were observed for catch per unit effort of Kamchatka flounder (Atheresthes evermanni), Pacific cod (Gadus macrocephalus), and walleye pollock, implying an important impact of predation by these groundfishes on JPS recruitment. The overall model incorporating these abiotic and biotic factors successfully reproduced the variability in JPS recruitment. Temperature and wind conditions around the spawning ground along with predator condition in the nursery ground appear to play a dominant role in the recruitment dynamics of JPS. Based on these results and prior knowledge, we propose a new hypothesis to explain the processes controlling JPS recruitment.     

Mortality of larval walleye pollock Theragra chalcogramma in the western Gulf of Alaska, 1988–91

Mortality rates of larval walleye pollock Theragra chakogramma were estimated from larval survey data from 1988 to 1991. Mortality estimates were based on cohort-specific losses between occupations of survey grids. Interannually, estimates of early feeding stage larval mortality rates ranged over an order of magnitude, from 0.045–0.43 day^-1, and declined sharply with age. There is some evidence that mortality rates of early feeding larvae tend to be negatively correlated with temperature and postively correlated with wind mixing.     

Predation on walleye pollock (Theragra chakograrnma) eggs and yolk-sac larvae by pelagic crustacean invertebrates in the western Gulf of Alaska

Immunological detection of yolk protein was used to assess predation by pelagic amphipods (gammarid and hyperiid), mysids, and euphausiids on eggs and yolk-sac larvae of walleye pollock Theragra chalcogramma during 1988 and 1989. Consumption estimates were made on the basis of frequency of positive immunoassays, assay detection times (gut clearance time), predator abundance, and spatial overlap of predators and prey. From our results gammarid amphipods and euphausiids were important predators on eggs and yolk-sac larvae, respectively. Gammarid amphipods alone consumed about 14% of the standing stock of pollock eggs in 1989. These results were compared with those from clearance rate experiments of predators feeding on pollock eggs in 300-1 bags. In general, clearance rate estimates of egg consumption were lower than those determined from gut contents.     

Early- to late-summer population growth and prey consumption by age-0 pollock (Theragra chalcogramma), in two years of contrasting pollock abundance near the Pribilof Islands, Bering Sea

Acoustic survey data were used to estimate the abundance and distribution of age-0 walleye pollock and zooplankton near the Pribilof Islands, Bering Sea, nursery area at two time periods in two consecutive years: the beginning of August, and mid-September, of 1996 and 1997. The 1996 pollock year class ultimately produced a large adult cohort in the eastern Bering Sea, while the 1997 year class produced a below-average adult cohort. Acoustic densities of age-0 pollock were significantly lower in August – and declined more strongly from August to September – in 1997 than in 1996, indicating that the trend to adult cohort strength was already set by August. Diet composition analyses revealed that age-0 pollock ate a much higher proportion of euphausiids in 1997 than in 1996, despite lower acoustic abundance of euphausiids in 1997. We infer that in 1996, age-0 pollock experienced greater feeding success by August, with high concentrations of copepods available for smaller fish to consume, and high concentrations of euphausiids available for larger individuals. In 1997, age-0 pollock had lower body condition in August and may have been limited by the availability of small (<2 mm) copepods. Bioenergetic modeling of prey consumption did not indicate a likelihood that age-0 pollock would begin to deplete euphausiids until late August in 1996, and not at all between August and mid-September in 1997.     

Interannual variability in growth of walleye pollock, Theragra chalcogramma, in the central Bering Sea

Interannual variability in growth of walleye pollock, Theragra chalcogramma, was examined. Adult walleye pollock were collected from the central Bering Sea (Aleutian Basin) from 1978 to 1999. Average fork lengths were found to be approximately 47 cm during the 1970–80s, this increased to 56 cm in the late 1990s. Age was determined for 4805 individuals using the otolith break and burn method. Ages ranged from 5–23 years and the year classes of 1978 and 1989 were dominant in the 1980s and the 1990s, respectively. Fish had significantly larger length-at-age in the 1990s compared to the 1970–80s, and interannual variability in age–length relationship was clearly observed. Taking into consideration a recent decrease of the walleye pollock biomass in the central Bering Sea, density-dependent growth was supported as one possibility of the growth variability. At the same time, we could not rule out the possibility that oceanographic variability affected the growth of walleye pollock in the area.     

Interannual variability in growth of larval and juvenile walleye pollock Theragra chalcogramma in the western Gulf of Alaska, 1983–91

Interannual variability in growth of larval walleye pollock Theragra chalcogramma was examined from 1983 to 1991 and of juveniles from 1985 to 1990. ANCOVA was used to assess differences in population growth rates, and an alternate method was developed to examine variations between annual length-at-age data and average 'expected' values over different age groupings. For larvae, the years 1986, 1987, 1989 and 1990 had higher than average length-at-age, and 1988 and 1991 had lower than average values. Relationships between growth and SST and larval density were not clear. A tentative relationship between copepod nauplii abundance and larval length-at-age was noted. The consequence of larval growth to larval mortality, late larval abundance or recruitment was not clear. We conclude that larval mortality rates are highly variable and tend to mask effects of moderate variability in growth on later abundance. For juveniles, 1987 had significantly lower than average length-at-age and 1988 had higher than average values. Although there are few years of data, they tend to support the importance of juvenile growth in the recruitment process. Conditions for the large 1988 year class are documented and discussed, including warm SST, calm winds, relatively low larval growth rates, low abundances of potential predators on larvae, low larval mortality rates, and high juvenile growth rates.     

Cohort survival patterns of walleye pollock, Theragra chalcogramma, in Shelikof Strait, Alaska: a critical factor analysis

A series of age-specific life tables for walleye pollock ( Theragra chalcogramma ) in the western Gulf of Alaska was compiled for the 1980-91 year classes. The life tables were utilized to perform an exploratory key factor analysis to examine the timing of critical periods in the recruitment process, evidence of density-dependence at different stages and trends in mortality rates. Early larval mortality was significantly correlated with generational mortality (In recruits/spawning bio-mass), but patterns in juvenile mortality also were similar to generational mortality and in some years were clearly dominant in determining the fate of a cohort. Density-dependent mortality, based on the correlation between mortality and initial abundance, was indicated only for the late larval to early juvenile stage. Time trends were marginally significant for juvenile mortality. It is speculated that the observed increase in juvenile mortality is associated with increasing abundance of arrowtooth flounder. Weaknesses in the data base are discussed; these along with the short time series involved make our conclusions tentative and subject to further study. We hypothesize that pollock recruitment levels can be established at any life stage depending on sufficient supply from prior stages, a type of dynamics which can be termed supply dependent multiple life stage control.     

Utilization of free amino acids, yolk proteins and lipids in developing eggs and yolk-sac larvae of walleye pollock Theragra chalcogramma

ABSTRACT:   To elucidate the utilization of the major yolk nutrient stocks in eggs and larvae of walleye pollock Theragra chalcogramma , the contents of free amino acids (FAA), the major yolk protein (180 kDa lipovitellin originated from vitellogenin B in ovulated eggs: oLv B), and lipids were measured. Most eggs hatched 18 days after fertilization at 5°C, and all larvae absorbed almost all their yolk mass by 28 days. The total FAA content showed no change during the first 6 days, and then decreased to 28% of the initial level by 18 days. The oLv B contents, measured by an enzyme-linked immunosorbent assay using a specific antiserum against oLv B, gradually decreased from 6 to 18 days, followed by a rapid decline. The content of phospholipids (PL) and triacylglycerols (TG) showed no marked change until hatching, and then decreased until disappearance of yolk sac. From these results, it is proposed that there are two main periods for nutrient utilization in embryos and larvae of walleye pollock. In the first period, FAA was mainly utilized until 18 days after fertilization. Active utilization of oLv B and lipids (PL and TG) instead of FAA occurred during the second period from 18 to 28 days.     

The influence of pelagic habitat selection and interspecific competition on productivity of juvenile walleye pollock (Theragra chalcogramma) and capelin (Mallotus villosus) in the Gulf of Alaska

Here we investigate processes affecting productivity of capelin and walleye pollock in the Gulf of Alaska. We examine pelagic habitat selection by comparing the distribution of juvenile fish and their prey with oceanographic properties and we evaluate the potential for interspecific competition by comparing diets and measures of foraging. The primary field study was conducted in Barnabus Trough, Kodiak Island, Alaska, during September 2005. The distribution of fish was assessed acoustically and trawls were used to collect individual fish for stomach content analyses. Physical and biological data were collected with conductivity–temperature–depth probes and zooplankton tows. Age‐0 pollock were distributed in cool waters offshore of a mid‐trough front, coincident with the distribution of euphausiids, their preferred prey. In contrast, capelin and their prey (copepods) were distributed throughout the trough. We observed that sympatric capelin (occurring with pollock) often had reduced foraging success compared to allopatric capelin (occurring alone). Results of a bioenergetic model also suggest that the exclusion of capelin from foraging on euphausiids can have negative consequences for capelin growth.     

A review of the distribution, ecology and population dynamics of age-0 walleye pollock in the Gulf of Alaska

This review paper synthesizes published research and unpublished data on the abundance and distribution patterns, ecology and population dynamics of walleye pollock ( Theragra chalcogramma ) during their first year of life (age-0) in the Gulf of Alaska. Distribution patterns have been described using mainly trawl catches, but recently, acoustic methodology has been employed, especially in examining vertical distributions. Although age-0 pollock are found throughout the Gulf, the highest catches occurred west of Kodiak Island. Pollock are pelagic for at least their first 6 months of life but show an ontogenetic increase in depth distribution superimposed on a pronounced diel vertical migration at a larger size. Daily growth rates are variable depending on year, season and area, and growth generally ceases during the winter. The diet of age-0 pollock shifts from mainly copepods in early juveniles to euphausiids by fall, with epibenthic organisms becoming important during the winter months. Feeding occurs mainly at night in surface waters. Age-0 pollock are most frequently associated with gelatinous zooplankton (medusae) and older pollock. Many predators on age-0 pollock have been identified; the most important are arrowtooth flounder ( Atheresthes stomias ), adult pollock, puffins ( Fratercula spp.), murres ( Una spp.), harbor seals ( Phoca vitulina richardsi ) and Steller sea lions ( Eumetopiasjubatus ). Modelling provides some insight into the population dynamics of these juveniles and environmental conditions which interannually affect their survival. These results are discussed relative to the importance of age-0 pollock in the recruitment of this species and to their role in the pelagic ecosystem.     

Physical transport of young pollock larvae (Theragra chalcogramma) near Shelikof Strait as inferred from a hydrodynamic model

An advective model was used to simulate the drift of larval walleye pollock ( Theragra chalcogramma ) over a 40-day period (late April through early June) near Shelikof Strait, Alaska. This model was used: (i) to assess how much of the observed change in larval positions during that period can be explained by transport at fixed depth; (ii) to demonstrate that observed change can be related to mean large-scale meteorological forcing; and (iii) to investigate accumulation of larvae in specific areas near the coast. Based on availability of larval and circulation data, three years were studied: 1988, 1989 and 1991. Velocity fields generated from a hydrodynamic model driven by winds and runoff were used to advect particles seeded in accordance with observed larval distributions in late April of each year. The modelled larvae were tracked at 40 m depth, corresponding to the mean depth of sampled larvae and the depth of neutrally buoyant drifters employed in field studies. Specific features observed in late May larval surveys were reproduced by the model, such as the accumulation of larvae in a shoal area downstream of the strait. Differences among the modelled years include extensive flushing of larvae to the south-west in 1988 and 1991, vs. limited flushing in 1989. These differences appear related to the mean large-scale atmospheric pressure patterns for April-May of those years.