Heterotrophic bacterial dynamics in the lower Fraser River,its estuary and Georgia Strait,British Columbia,Canada

Annual cycles in the glucose heterotrophic potentials of the bacterioplankton of the lower Fraser River, its estuary and Georgia Strait surface waters were observed. Maximal microbial heterotrophic potentials were noted during the winter and early spring months, whereas maximal heterotrophic productivities were observed to occur during the summer and fall months. Temperature and soluble organic carbon levels appear to be the most important of the assayed physico-chemical parameters in regulating activities and productivities of the heterotrophic microbes of these waters.     

Simulation study of phytoplankton photosynthesis and growth in the Fraser River estuary

Phytoplankton photosynthesis was estimated by a simulation model of the Fraser River estuary (Canada). For this estimation, two kinds of information were considered; (1) the photosynthetic response of phytoplankton taken from the estuary to each environmental factor; (2) actual measurement of changes in each environmental factor in the estuary. As the major limiting factors, light intensity, temperature, and nitrate were taken into account. Variations in these parameters were combined together under the assumption of the law of the minimum; that is, only one of the environmental factors was considered to limit photosynthesis at any given time and place. From the results obtained, it could be shown which environmental factors actually limited phytoplankton photosynthesis in the Fraser River estuary at different times of the year and at different depths.     

Heterotrophic activity associated with particulate size fractions in a Spartina alterniflora salt-marsh estuary,Sapelo Island,Georgia, USA,and the continental shelf waters

An investigation was undertaken to determine the distribution and activity of the heterotrophic microplankton associated with particles from a Georgia salt marshestuarine ecosystem to the western edge of the Sargasso Sea. Heterotrophic activity was determined by the uptake of ¹⁴C-glucose. More than 80% of the activity was associated with detritus greater than 3 m in creek, river and coastal (within 4 km of shore) waters. In the Gulf Stream, approximately 80% of the heterotrophic activity was in the fraction less than 3 m. In the estuary, total heterotrophic activity fluctuated with the tides; the greatest activity occurred near low ebb tide at all locations. The lowest activity was measured at slack low and high tides. In creek water most of the heterotrophic activity was associated with particles between 14 and 180 m, whereas in coastal waters (less than 4 km from shore) most of the activity was in the fraction greater than 180 m.Contribution No. 331 from the University of Georgia Marine Institute, Sapelo Island, Georgia 31327, USA.     

Shallow-water benthic and pelagic metabolism:

Pelagic primary production and benthic and pelagic aerobic metabolism were measured monthly at one site in the estuarine plume region of the nearshore continental shelf in the Georgia Bight. Benthic and water-column oxygen uptake were routinely measured and supplemented with seasonal measures of total carbon dioxide flux. Average respiratory quotients were 1.18:1 and 1.02:1 for the benthos and water column, respectively. Benthic oxygen uptake ranged from 1.23 to 3.41 g O₂ m^-2 d^-1 and totalled 756 g O₂ m^-2 over an annual period. Water column respiration accounted for 60% of total system metabolism. Turnover rates of organic carbon in sediment and the water column were 0.09 to 0.18 yr^-1 and 6.2 yr^-1, respectively. Resuspension appeared to control the relative amounts of organic carbon, as well as the sites and rates of organic matter degradation in the benthos and water column. Most of the seasonal variation in benthic and pelagic respiration could be explained primarily by temperature and secondarily by primary productivity. On an annual basis, the shelf ecosystem appeared to be heterotrophic; primary production was 73% of community metabolism, which was 749 g C m^-2 yr^-1. The timing of heterotrophic periods through the year appeared to be closely related to both river discharge and the periodicity of growth and death of marsh macrophytes in the adjacent estuary. The results of this study support the estuarine outwelling hypothesis of Odum (1968).This is Contribution No. 530 from the University of Georgia Marine Institute. This work was supported by the Georgia Sea Grant College Program maintained by the National Oceanic and Atmospheric Administration, US Department of Commerce     

Behavior of a Small Pulsed River Plume in a Strong Tidal Cross-Flow in the Akashi Strait

We carried out a field study of a small river plume emptying into Osaka Bay near the Akashi Strait in western Japan, to understand the fate of its urban-runoff-laden waters. As the river is weak compared to tidal currents, we find that the behavior of the plume is strongly influenced by phasing between tidal stage and currents (a westward-traveling progressive tidal wave). When the tidal stage rises, sea water floods up the river, and concentrated river effluent cannot be seen in Osaka Bay. During most of the river’s ebb, a topographic eddy transports river effluent towards the energetic Akashi Strait, while strong vertical shear in the tidal flow mixes the effluent with seawater. However, there is a short interval of time during which the direction of tidal flow in the Strait changes direction and the magnitude of the current is weak. At this time, shear is weak enough to allow a stratified plume of concentrated river water to form, and this plume is driven offshore, and into Osaka Bay, by baroclinic circulation. A sewage outfall, which is located very close to the river mouth, is subjected to these same conditions and therefore exhibits similar behavior. Understanding the relation between tidal phase and plume behavior will be useful to Kobe City, as well as other cities in a similar environment, for minimizing the environmental effects of their wastewater and non-point-source runoff.     

Dependence of dissolved carbohydrate concentrations upon small scale nanoplankton and bacterioplankton distributions in the western Sargasso Sea

Significant correlations between trophic components of the <20 m microbial plankton and the concentrations of total dissolved carbohydrate (TCHO) and polysaccharide (PCHO) have been found at two drogued buoy stations which were sampled at 3 or 4-h intervals over diel cycles. An attempt was made to sample on two pre-selected isotherms (approximating isopycnals) at each station. Significant inverse correlations of TCHO and of PCHO with numbers of phototrophic nanoplankton (PNAN) were found along both isotherms at a station off the Carolina coast. Off central Florida, TCHO and/or PCHO were significant multiple linear functions of PNAN, heterotrophic bacterioplankton (BAC) and sometimes heterotrophic nanoplankton (HNAN) counts. Partial regression coefficients for PNAN were always negative and those for HNAN were positive. Inverse trends dominated the relations of BAC with PCHO and TCHO. The combined data from the two stations (300 nautical miles apart) produced very similar multiple linear relations. This suggests that the observed relationships are real, resulting from general physiological processes and interactions of the microbial plankton groups rather than from chance occurrences at a particular station, and that the combined activities of the <20 m plankton actively regulate dissolved carbohydrate concentrations in the Sargasso Sea.     

Some experiments on phosphate assimilation by coastal marine plankton

A study of phosphate assimilation by coastal marine plankton revealed that both phytoplankton and microheterotrophs incorporated radioactive phosphorus (³³P). Size fractionation of the particulate matter (using 1 m pore diameter Nucleopore^® membrane filters), antibiotic treatment (using garamycin), and independent estimaties of photoautotrophic (¹⁴CO₂ uptake) and heterotrophic (³H-glucose uptake) activities were employed to separate phyto- and bacterioplankton phosphate uptake. Results indicated that phytoplankton ³³P-uptake was best estimated by the fraction of particulate matter retained on the 1 m membrane filters. Usually, less than 10% of the phytoplankton (based on chlorophyll a measurements) passed the 1 m pore-diameter filters, whereas about 90% of the heterotrophic activity passed. At least 50% of the ³³P-uptake was associated with the <1 m fraction. It may be possible to resolve the phytoplankton and bacterial contributions to ³³P-uptake by comparing the percent of total ³³P-uptake with the percent of total ³H-glucose uptake associated with the >1 m fraction.     

Thymidine incorporation as a measure of heterotrophic bacterioplankton production in marine surface waters: Evaluation and field results

To assess bacterioplankton production in the sea, we have developed a procedure for measuring growth based on incorporation of tritiated thymidine into DNA; the accuracy of this procedure was tested under a variety of laboratory and field conditions. By autoradiography, we have found that for all practical purposes our technique is specific for the nonphotosynthetic bacteria and that virtually all of the active bacteria (one-third or more of the total countable bacteria) take up thymidine. We also measured (1) the intracellular isotope dilution of thymidine assessed by parallel experiments with labeled phosphorus, and (2) DNA content of natural marine bacteria (0.2 to 0.6 m size fraction); a conversion factor derived from these data permitted estimation of production from thymidine incorporation results. A very similar conversion factor was independently derived from the empirical relationship between thymidine incorporation and growth of natural bacterioplankton under controlled conditions. Combined results show that this technique, which can be performed rapidly and easily at sea, provides good estimates of production. Data from Southern California Bight waters, which contain oligotrophic as well as moderately eutrophic regions, show that average bacterioplankton doubling times, like those of the phytoplankton, are on the order of a few days, with fastest growth at depths just below those of greatest phytoplankton abundance. Offshore bacterial production is roughly 5 to 25% of the primary production; thus, at a 50% assimilation efficiency, the bacterioplankton would consume 10 to 50% of the total fixed carbon.     

Feeding in a larval fish assemblage: the nutritional significance of an estuarine plume front

Gut fullness of larval fishes was used to determine the nutritional significance of an estuarine plume front off Botany Bay, SE Australia, on three days in March/April 1990. Fishes were captured in three different water masses (estuarine plume, front and shelf water), each separated by 200 m, using a 260 m mesh purse-seine net. Overall, the gut-fullness index (GFI) of 260 fish larvae combined from eight families (Gerreidae, Mugilidae, Mullidae, Sparidae, Blenniidae, Kyphosidae, Monodactylidae, Pomacentridae), was significantly greater in the plume and front water on two of the sampling occasions, but no difference was detected on the third occasion. Trends in GFI among families were inconsistent with respect to the front. The mugilid (Liza argentea) fed equally and abundantly in all water masses (gut fullness>90%), while the kyphosid (Kyphosus spp.) had a significantly greater GFI in the plume compared to the shelf water. In general, the response of GFI to the front varied between dates and amongst taxa. Diet analysis showed that mugilids selected for copepod nauplii within the plume and shelf water [alpha selectivity index (ASI) of (0.22 to 0.98)] and for harpacticoid copepods within the front (ASI=0.92), representing characteristic diets identified by canonical discriminant analysis. Kyphosids selected positively for copepod nauplii in all three water masses on both occasions (ASI>0.96). In general, diets of larval fish were taxon-specific, and responded variably to the three habitats. Comparison of diets across water masses was complicated by the large number of families, which rarely occurred in all water masses. The nutritional significance of an estuarine plume front varies between species of larval fish, and there are no obvious trends that can be applied to the larval fish community in general.     

Numerical analysis on the Brisbane River plume in Moreton Bay due to Queensland floods 2010–2011

During the Queensland floods in the summer of 2010–2011, a flood-driven Brisbane River plume extended into Moreton Bay, Queensland, Australia, and then seaward, travelling in a northward direction. It covered approximately 500 km $^{2}$ . This paper presents a three- dimensional hydrodynamic numerical model investigation into the behaviour of the Brisbane River plume. The model was verified by using satellite observations and field measurement data. The present study concludes that the high river discharge was the primary factor determining the plume size and its seaward extensions. A notable finding was that the plume was a bottom-trapped type rather than a buoyant type. Further, the southerly winds were found to have moderately confined the alongshore extension of the plume, and had caused the plume to mix thoroughly with the ocean water.     

Efficient size-selective bacterivory by phagotrophic nanoflagellates in aquatic ecosystems

Size-selective grazing on bacterioplankton by phagotrophic nanoflagellates was analyzed and modelled. The proposed model resembles a Monod equation (Michaelis-Menten-like expression) in which clearance rates by heterotrophic nanoflagellates depend on bacterioplnnkton biovolume. Larger bacteria were ingested faster than smaller bacteria. The proposed model was in agreement with experimental data from different authors, both from cultures and natural assemblages. Size-selective grazing efficiency was analyzed as the ability of phagotrophic nanoflagellates to discriminate between cells differing in volume by a factor of two, e.g., corresponding to dividing and nondividing bacteria. Unlike previously published models, the proposed model suggests that phagotrophic nanoflagellates could be highly effective size-selective grazers for small bacteria (<0.1 m³), which are the most common bacterial sizes in planktonic systems. However, phagotrophic nanoflagellates were unable to size-discriminate dividing and nondividing bacteria for volumes >0.1 m³. These results strongly support the hypothesis that heterotrophic nanoflagellate grazing, by discriminating between different sizes of dividing and nondividing bacteria, may actually be regulating bacterial size and growth rate in natural aquatic ecosystems.     

厦门后溪流域沿城乡梯度浮游细菌多样性及其与环境因子的关系

为研究城市化对河流浮游细菌的影响,选取厦门后溪流域沿城乡梯度采集样品,并利用变性梯度凝胶电泳(Denaturing gradient gel electrophoresis,DGGE)和多元统计方法对浮游细菌群落空间分布格局进行分析.结果表明,Shannon-Weiner多样性指数和DGGE条带数随着采样站点和城市中心距离的减小呈现先升高后下降的趋势.聚类分析(Cluster)和多维尺度分析(Multidimensional scaling,MDS)将12个站点的浮游细菌群落分为4组,测序结果显示β-变形菌(Betaproteobacteria)是优势类群,占40.0%.统计分析表明,pH和总氮(Total nitrogen,TN)是影响后溪浮游细菌群落组成最主要的环境因子,他们共同解释了47.6%的群落组成变化.总之,沿城乡梯度随着水体理化指标的变化,浮游细菌群落也发生明显转变,表明后溪浮游细菌受到城市化的强烈影响.此外,结果也提示DGGE可以作为监测与评价河流生态系统健康的重要工具.图6表2参33     

A comparison of metals in sediments and water in the River Nahr-Ibrahim,Lebanon: 1996 and 1999

Rivers whose basins are underlain by carbonate rocks exhibit high pH, lower desorption of metals and possess high buffering capacity against acidic inputs to the river. The catchment of River Nahr-Ibrahim, Lebanon, is largely underlain by limestone. Compared to neighbouring countries, Lebanon is relatively fortunate since precipitation is high. However, recently a warming in temperature and a drop in precipitation has occurred, thus causing low water levels in rivers. The objective of this study is to investigate the variation of the total metal content (Fe, Mn, Zn, Cu, Pb and Cd) in bed sediments and water of River Nahr-Ibrahim between 1996 and 1999 (two years); and relate these variations to the effect of changes in human activities and/or due to the variations of precipitation rate, temperature and pH of water. Bed load sediments and stream water were collected simultaneously from five sampling sites. Water pH and temperature were determined in situ. Sediment samples were dried at room temperature and sieved; the sediment size <75 m size was retained. Water was analysed for major constituents and trace metals. Metals were extracted from sediments with aqua regia. Metal concentration in water and sediments were determined using ICP-MS technique. Data revealed a drop in metal concentrations (Zn, Cu, Pb, Cd) in sediments at quarry site after its closure. The decrease in precipitation rate, lowering the level of water and the dilution of industrial discharges and decrease in water pH led most probably to the desorption of metals from sediments into the water.     

A brominated secondary metabolite synthesized by the cyanobacterial symbiont of a marine sponge and accumulation of the crystalline metabolite in the sponge tissue

The dictyoceratid marine sponge Dysidea herbacea (Keller, 1889) is common in shallow waters of the tropical Pacific Ocean. Polybrominated biphenyl ethers such as 2-(2,4-dibromophenyl)-4,6-dibromophenol (1) are characteristic secondary metabolites of some specimens of this sponge and may represent as much as 12% of the dry weight. We have found 1 to be deposited as conspicuous crystals throughout the sponge tissue. The dominant prokaryotic endosymbiont in the mesohyl of the sponge is a filamentous cyanobacterium (Oscillatoria spongeliae), although a vacuole-containing, heterotrophic bacterium is also present. The cyanobacteria were separated from the sponge cells and heterotrophic bacteria by flow cytometry. Coupled gas chromatography—mass spectrometry and proton nuclear magnetic-resonance spectroscopy revealed that the major brominated Compound 1 isolated from the intact symbiotic association is found in the cyanobacteria and not in the sponge cells or heterotrophic bacteria. This suggests that the production of the compound is due to the cyanobacterium, and not to the sponge or symbiotic heterotrophic bacteria, as had been suggested earlier.     

The trophic role of glycolic acid in coastal seawater. I. Heterotrophic metabolism in seawater and bacterial cultures

Glycolic acid, a known algal excretory product, represents a potentially important energy source for heterotrophic bacteria in marine waters. Measurements of heterotrophic uptake and mineralization by the natural microbes in the plankton indicate quantitative use of glycolic acid comparable to other common microbial substrates. This activity shows vertical and horizontal variations that correlate in a general way with primary productivity. Glycolic acid appears to be present at higher concentrations than other low molecular weight substrates, and it also shows a lower rate of turnover. It is mineralized (respired) to a greater extent than other recorded substrates, an average of 70% of total uptake. Two-thirds of a total of 141 colonies of bacteria cultured from seawater on marine agar proved capable oftaking up and respiring glycolic acid. These same bacteria, however, were unable to utilize glycolic acid for growth when it was the sole carbon source. Glycolate may therefore occur at higher concentrations and show erratic fluctuations in natural waters because microbial use is not tied to glycolate production but to the presence of other substrates. The fact that so many bacteria are able to metabolize it indicates a possibly important trophic role as an energy source. It is suggested that glycolic acid may be a major source of energy for active transport of other substrates by marine bacteria.     

Integral models for bubble,droplet, and multiphase plume dynamics in stratification and crossflow

We present the development and validation of a numerical modeling suite for bubble and droplet dynamics of multiphase plumes in the environment. This modeling suite includes real-fluid equations of state, Lagrangian particle tracking, and two different integral plume models: an Eulerian model for a double-plume integral model in quiescent stratification and a Lagrangian integral model for multiphase plumes in stratified crossflows. Here, we report a particle tracking algorithm for dispersed-phase particles within the Lagrangian integral plume model and a comprehensive validation of the Lagrangian plume model for single- and multiphase buoyant jets. The model utilizes literature values for all entrainment and spreading coefficients and has one remaining calibration parameter \( \kappa \), which reduces the buoyant force of dispersed phase particles as they approach the edge of a Lagrangian plume element, eventually separating from the plume as it bends over in a crossflow. We report the calibrated form \( \kappa = [(b - r) / b]^4 \), where b is the plume half-width, and r is the distance of a particle from the plume centerline. We apply the validated modeling suite to simulate two test cases of a subsea oil well blowout in a stratification-dominated crossflow. These tests confirm that errors from overlapping plume elements in the Lagrangian integral model during intrusion formation for a weak crossflow are negligible for predicting intrusion depth and the fate of oil droplets in the plume. The Lagrangian integral model has the added advantages of being able to account for entrainment from an arbitrary crossflow, predict the intrusion of small gas bubbles and oil droplets when appropriate, and track the pathways of individual bubbles and droplets after they separate from the main plume or intrusion layer.     

Identification of Amazon River water at Barbados,W. Indies,by salinity and silicate measurements

Salinity and silicate concentrations were studied at about fortnightly intervals for 21 months at a station near Barbados, W. Indies; latitude 13°15 N, longitude 59°42 W. A sensitive inverse correlation was found to exist at 5 and 25 m, but not at greater depths. Salinity near the surface varied between 33.5 and 36.0, and silicate between a little less than 1 and 4 g at/l. Low salinity water, rich in silicate, was found from February to July; salinity increased and silicate decreased from September to December. It is argued that the low salinity water at Barbados can be identified with the areas of reduced salinity found by Ryther et al. (1967) about latitude 8° to 10° N, longitude 50° to 55° W, and that this water originates from the Amazon River. Local precipitation does not seem to be a significant factor.     

Distribution and structure of heterotrophic protist communities in the northeast equatorial Pacific Ocean

The distribution and structure of heterotrophic protist communities and size-fractionated chlorophyll a were studied during the Korea Deep Ocean Study 98 (KODOS 98) research expedition (July 1998) in the northeast equatorial Pacific Ocean (5–11°N). Areas of convergence and divergence formed at the boundaries of the South Equatorial Current (SEC), North Equatorial Current (NEC), and North Equatorial Counter Current (NECC) during the expedition. Water column physicochemical characteristics significantly influenced the size structure of heterotrophic protist communities. Intense vertical mixing and high nutrient and chlorophyll a concentrations characterized SEC and NECC areas, which were affected by converging and diverging water masses, respectively. Nanophytoplankton dominated in SEC and NECC areas; both areas also had relatively high heterotrophic protist biomasses (average 743 µg C m^–2). NEC areas were characterized by a stratified vertical structure, low nutrient and chlorophyll a concentrations, and picophytoplankton dominance. The heterotrophic protist biomass in NEC areas averaged 414 µg C m^–2; nanoprotists (<20 µm) dominated the community. The nanoprotist biomass comprised 49–54% of the total heterotrophic protist biomass in SEC/NECC areas and 67–72% in NEC areas. The biomass of heterotrophic protists was higher in SEC/NECC areas than in NEC areas, but the relative importance of nanoprotists was greater in NEC areas than in SEC/NECC areas. Heterotrophic dinoflagellates were dominant components of the <20 µm and >20 µm size classes in both water columns. The biomass of heterotrophic protists significantly correlated with the net-, nano-, and picophytoplankton biomass in SEC/NECC areas and with the nano- and picophytoplankton biomass in NEC areas. Heterotrophic protists and phytoplankton also showed strong positive correlation in the study area. The size structure of the phytoplankton biomass coincided with that of heterotrophic protists; the heterotrophic protist biomass positively correlated with the protists prey source. These relationships suggest that the community structure of heterotrophic protists and the microbial food web depended on size classes within the phytoplankton biomass. Microzooplankton grazing and phytoplankton growth rates were higher in SEC/NECC areas than in NEC areas. In contrast, the potential primary production grazed by microzooplankton was relatively high in NEC areas (127.3%) compared with SEC/NECC areas (94.6%). Our results indicate that the relative importance and size structure of heterotrophic protists might vary according to two distinct water column structures.Communicated by T. Ikeda, Hakodate     

Determination and properties of actively metabolizing heterotrophic bacteria in the sea,investigated by means of micro-autoradiography

Substrate transformation and microbial biomass production in aquatic ecosystems depend mainly on the total number of actively metabolizing heterotrophic bacteria. The most common methods used concern the determination of either the colony-forming bacteria or the total number of bacteria including autotrophs and inactive organisms a micro-autoradiographic method is presented which enables the substrate uptake of single bacteria by means of ³H-amino-acid mixture and Nuclepore filters to be determined. The standardization procedure revealed the greatest success after 3 h incubation with 10 Ci/ml tritiated amino-acid mixture and an exposure of 14 days to the X-ray film. Preliminary experiments showed inactivation of an active fresh-water population from 100% to 0.6% within 3 h at 28S. With increasing distance from the shore, the number of colony-forming units decreases from 6 to 0.01% of the total number of active heterotrophic bacteria. It is concluded from the results that the fraction of very small heterotrophic bacteria which cannot be cultured on nutrient media is responsible for the continuous breakdown of organic matter in off-shore regions of the sea.     

Evidence for multiple recruitment-cohorts of Puget Sound Dungeness crab,Cancer magister

While sampling intertidally in Puget Sound, Washington, USA, for juvenile Dungeness crab (Cancer magister) in 1984, we found evidence of two distinct cohorts of the same year-class based on sizes of first-instar juveniles (J1) and the spatial/temporal patterns of settlement. In 1988, three distinct cohorts were observed to settle in Puget Sound and its approaches. Settlement of one cohort occurred during May in the Strait of Juan de Fuca and in those areas of Puget Sound closest to the Strait. J1 individuals of this cohort were large (x=7.4 mm carapace width, CW) and comparable in both size and timing of settlement to populations along the Washington coast (e.g. Grays Harbor and Willapa Bay). Initial settlement density of the May cohort was as high as 215 crabs/m² in intertidal eelgrass beds along the Strait of Juan de Fuca and decreased to <2 crabs/m² within Puget Sound and the Strait of Georgia. A second cohort apparently originated in Hood Canal (a deep inland fjord), its size upon settlement in June was significantly smaller (J1 x= 5.3 mm CW) than the May cohort, and it was limited to Hood Canal and areas of Puget Sound close to the mouth of Hood Canal. A third cohort, which settled in late July and August, was the smallest of the three cohorts (J1 x= 4.8 mm CW), and was widely distributed around Puget Sound from Seattle in the south to the USA/Canadian border in the north. We hypothesize that most juvenile recruitment in Hood Canal and Puget Sound originates from parental stocks endemic to their respective basins (Hood Canal and Puget Sound cohorts), but that, on occasion, oceanographic conditions allow substantial influx of Pacific Ocean Dungeness crab larvae (oceanic cohort) through the Strait of Juan de Fuca into Puget Sound. Tracking of spatial/temporal settlement patterns and comparison of J1 sizes proved useful for estimating the probable sources and dispersion of Dungeness crab larvae. Differences in size and time of settlement between various larval cohorts of C. magister may prove useful as biomarkers for tracing circulation patterns within and between inland waters of Washington and the Pacific Ocean. Causes of smaller size and later settlement of the Puget Sound cohort relative to oceanic conspecifics of the same year-class are discussed.Contribution No. 856, School of Fisheries, University of Washington, Seattle