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Sedimentary phosphorus cycling and a phosphorus mass balance for the Green Bay (Lake Michigan) ecosystem

The tributaries of Green Bay have long been recognized as major sources of phosphorus in the Lake Michigan basin. The status of Green Bay as a sink or source of phosphorus for Lake Michigan proper has been less well defined. The bay receives nearly 70% of its annual load of phosphorus ( 700 metric tons (t) &middot; year-1) from a single source: the Fox River. Most of this phosphorus is deposited in sediments accumulating at rates that reach 160 mg &middot; cm-2 &middot; year-1 with an average of 20 mg &middot; cm-2 &middot; year-1. The phosphorus content of these sediments varies from <5 to >70 &micro;mol &middot; g-1. Deposition is highly focused, with ~70% of the total sediment accumulation and at least 80% of the phosphorus burial occurring within 20% of the surface area of the bay. Diagenetic and stoichiometric models of phosphorus cycling imply that >80% of the phosphorus deposited is permanently buried. External phosphorus loading to the bay is combined with sediment fluxes of phophorus to arrive at a simple phosphorus budget. Green Bay acts as an efficient nutrient trap, with the sediments retaining an estimated 70-90% of the external phosphorus inputs before flowing into Lake Michigan.

Wisconsin

Vertical distribution of the chloromonad flagellate Heterosigma carterae in columns: Implications for bloom development

Blooms of the marine flagellate Heterosigma carterae have been associated with catastrophic fish kills at mariculture facilities around the world. The precise cause(s) of the sudden appearance and disappearance of Heterosigma surface blooms has not been completely described or understood. Environmental data from prior studies of blooms indicate the presence of vertical stratification of the water column that is often induced by freshwater runoff. We report the relatively rapid concentration of Heterosigma cells at the surface of tubes shortly after the addition of distilled water to the surface. This phenomenon of cell concentration in surface waters may partially explain the sudden appearance of Heterosigma cells near the surface and subsequent disappearance when vertical stratification is lost due to turbulence or mixing. The results may lead to bloom mitigation techniques for net pen aquaculture.

Canadian Journal of Fisheries and Aquatic Sciences

The loss of hyperosmoregulatory ability in migrating juvenile American shad, Alosa sapidissima

Investigations on juvenile American shad ( Alosa sapidissima ) revealed several physiological changes associated with downstream migration. Plasma chloride decreased 20% in wild juvenile shad during the autumn migration. Migrants had lower condition factor and hematocrit than non-migrant shad captured by beach seining. Gill Na + ,K + -ATPase activity of migrant shad was higher than non-migrant; a 2.5-fold increase was observed in 1993, while a 57% increase was observed in 1994. Similar changes were observed in laboratory studies of shad maintained in fresh water under simulated natural temperature and photoperiod. Plasma chloride dropped 68% and gill Na + ,K + -ATPase activity increased 3-fold over a 3-month period. Decreased plasma chloride was associated with increased mortality. Increases in gill Na + ,K + -ATPase activity decreases in plasma chloride and osmolality, and incidence of mortality were delayed and moderated, but not eliminated, in shad maintained at constant temperature (24°C). Shad did not survive in fresh water past December regardless of temperature regime. In seawater, all shad survived and showed no perturbation of plasma chloride at 24°C or simulated natural temperature (above 4°C). The decline in hyperosmoregulatory ability, as influenced by declining temperatures, may serve as a proximate cue for autumnal migration.

Canadian Journal of Fisheries and Aquatic Sciences

Gillnet selectivity for lake trout ( Salvelinus namaycush ) in Lake Superior

Gillnet selectivity for lake trout (Salvelinus namaycush) was estimated indirectly from catches in nets of 102-, 114-, 127-, 140-, and 152-mm stretch measure. Mesh selectivity was modeled as a nonlinear response surface that describes changes in the mean, standard deviation, and skewness of fish lengths across mesh sizes. Gillnet selectivity for lake trout was described by five parameters that explained 88% of the variation in wedged and entangled catches, 81% of the variation in wedged catches, and 82% of the variation in entangled catches. Combined catches of wedged and entangled lake trout were therefore described more parsimoniously than separate catches of wedged and entangled lake trout. Peak selectivity of wedged and entangled fish increased from 588 to 663 mm total length as mesh size increased from 102 to 152 mm, and relative selectivity peaked at a total length of 638 mm. The estimated lake trout population size-frequency indicated that gillnet catches were negatively biased toward both small and large lake trout. As a consequence of this bias, survival of Lake Superior lake trout across ages 9-11 was underestimated by about 20% when the catch curve was not adjusted for gillnet selectivity.

Canadian Journal of Fisheries and Aquatic Sciences

Characterization and application of a quantitative DNA marker that discriminates sex in Chinook salmon ( Oncorhynchus tshawytscha )

A qualitative male-specific DNA marker (OT-24) was amplified by spPCR (single-primer polymerase chain reaction) from chinook salmon (Oncorhynchus tshawytscha) DNA along with several non-sex-linked products. The termini of the male-specific product were sequenced, and a pair of PeR primers were constructed for marker-specific PCR amplification. Dual primer PCR (dpPCR), with the marker-specific primers, amplified a product from both nudes and females. The amount of dpPCR product amplified from males was at least 100-fold greater than that from females. The quantitative difference between males and females was consistent among geographically distinct populations from western U.S. rivers. In addition, DNA sequence analysis indicated that OT-24 was highly conserved among geographically distinct salmon populations. The qualitative spPCR product segregated through several genetic crosses indicating equal sex ratios among progeny. Identification of the male and female juveniles by dpPCR was consistent with the spPCR analysis. There was no tissue specificity observed by spPCR or dpPCR analysis of this marker. A rapid DNA extraction method and dpPCR analysis were used to nonlethally determine sex ratios in wild spring chinook salmon adults, withheld for genetic and behavioral studies, prior to their development of gross sexual differences in their external morphology.

Canadian Journal of Fisheries and Aquatic Sciences

Effects of surgically and gastrically implanted radio transmitters on swimming performance and predator avoidance of juvenile chinook salmon ( Oncorhynchus tshawytscha )

Radiotelemetry data are often used to make inferences about an entire study population; therefore, the transmitter attachment method should be the one that least affects the study animal. Juvenile chinook salmon ( Oncorhynchus tshawytscha ) <120 mm in fork length (FL) with either gastrically or surgically implanted transmitters had significantly lower critical swimming speeds than control fish 1 and 19-23 days after tagging. For fish >120 mm FL, fish with gastric implants swam as well as controls 1 day but not 19-23 days after tagging. In contrast, fish with surgical implants swam as well as controls 19-23 days but not 1 day after tagging. During predation trials, fish with gastric or surgical implants were eaten by smallmouth bass ( Micropterus dolomieu ) in significantly greater numbers than controls. We do not recommend implanting transmitters (representing 4.6-10.4% of the fish's body weight) in fish <120 mm FL. Furthermore, surgical implants (representing 2.2-5.6% of the fish's body weight) may be the preferred method for biotelemetry studies of juvenile chinook salmon >120 mm FL.

Canadian Journal of Fisheries and Aquatic Sciences

Methyl mercury dynamics in littoral sediments of a temperate seepage lake

The sites and rates of methyl mercury (MeHg) production and transport in littoral zone sediments were investigated at Pallette Lake in northern Wisconsin. In littoral areas where groundwater inflow occurs, sulfate supply from groundwater creates profiles of electron acceptors (sulfate) and donors (methane, sulfide) that are reversed from those found in sediments whose sulfate supply is delivered from overlying water. The highest MeHg concentrations in porewaters and the maximal advective MeHg flux rates (4.5-61.7 ng ·m -2 ·day -1 ) were observed in the spring, while highest bulk phase concentrations occur later in the summer. These estimated MeHg fluxes are greater than the mean areal production rates estimated previously for the water column and are similar to the atmospheric flux. Gross MeHg production was measured using the addition of 203 Hg as a tracer to sediments. The depth at which maximal 203 Hg methylation occurred coincided with the observed maximums in solid-phase and porewater MeHg concentrations. Because input, advection, and accumulation of MeHg in these sediments were measured directly, an independent estimate of MeHg production could be made and compared with 203 Hg-derived rates. This comparison suggests that that the 203 Hg tracer method provides reasonable estimates of gross methylation rates and that a substantial fraction of solid-phase Hg is available for methylation

Canadian Journal of Fisheries and Aquatic Sciences

Benthic invertebrate distributions in the San Joaquin River, California, in relation to physical and chemical factors

The invertebrate fauna of nontidal portions of the lower San Joaquin River and its major tributaries is described in relation to water quality and habitat using canonical correspondence analysis, autecological metrics, and indicator species analysis. A large-scale (basin-wide) pattern in community response to salinity (sulfate-bicarbonate type) was detected when standardized, stable substratum was sampled. Community structure, taxa richness, and EPT (ephemeropterans, plecopterans, and trichopterans) richness varied with dissolved solids concentration (55-1700 mg total dissolved solids. L-1), and distributions of many taxa indicated salinity optima. Distinct assemblages associated with either high or low salinity were evident over this range. Large-scale patterns in community structure were unrelated to pesticide distributions. Structure and taxa richness of invertebrate assemblages in sand substratum varied both with salinity and with microhabitat heterogeneity. The benthic fauna generally was dominated by a taxa-poor assemblage of specialized psammophilous species, contributing to a weaker relationship between community structure and water quality than was observed using standardized substratum. Habitat types and associated dominant species were characterized using indicator species analysis. Species assemblages did not vary substantially with irrigation regime or fiver discharge, indicating that structure of invertebrate communities was a conservative measure of water quality.

California

Competition between larval lake herring (Coregonus artedi) and lake whitefish (Coregonus clupeaformis) for zooplankton

Diet and growth of larval lake herring (Coregonus artedi) and lake whitefish (Coregonus clupeaformis) were compared in mesocosm experiments in a small mesotrophic lake in southeastern Michigan. Fish were sampled from single-species and mixed assemblages in 2-m 3 cages for 8 weeks during April and May. Both species initially ate mostly cyclopoid copepodites and small cladocerans (Bosmia spp.). Schoener's index of diet overlap showed considerable overlap (70-90%). Lake whitefish ate Daphnia spp. and adult copepods about 2 weeks earlier than did lake herring, perhaps related to their larger mean mouth gape. Lake whitefish were consistently larger than lake herring until the eighth week, especially in the sympatric treatments. Lake whitefish appeared to have a negative effect on the growth of lake herring, as lake herring in mixed-species treatments were smaller and weighed less than lake herring reared in single-species treatments. The diet similarities of lake whitefish and lake herring larvae could make them competitors for food in the Great Lakes. The greater initial size of lake whitefish could allow them to eat larger prey earlier and thereby limit availability of these prey to lake herring at a crucial period of development.

Michigan

Incubation temperature, developmental biology, and the divergence of sockeye salmon ( Oncorhynchus nerka ) within Lake Washington

Sockeye salmon ( Oncorhynchus nerka) introduced into Lake Washington in the 1930s and 1940s now spawn at several different sites and over a period of more than 3 months. To test for evolutionary divergence within this derived lineage, embryos that would have incubated in different habitats (Cedar River or Pleasure Point Beach) or at different times (October, November, or December in the Cedar River) were reared in the laboratory at 5, 9, and 12.5??C. Some developmental variation mirrored predictions of adaptive divergence: (i) survival at 12.5??C was highest for embryos most likely to experience such temperatures in the wild (Early Cedar), (ii) development rate was fastest for progeny of late spawners (Late Cedar), and (iii) yolk conversion efficiency was matched to natural incubation temperatures. These patterns likely had a genetic basis because they were observed in a common environment and could not be attributed to differences in egg size. The absolute magnitude of divergence in development rates was moderate (Late Cedar embryos emerged only 6 days earlier at 9??C) and some predictions regarding development rates were not supported. Nonetheless our results provide evidence of adaptive divergence in only 9-14 generations.

Washington

DNA and allozyme markers provide concordant estimates of population differentiation: Analyses of U.S. and Canadian populations of Yukon River fall-run chum salmon ( Oncorhynchus keta )

Although the number of genetic markers available for fisheries research has steadily increased in recent years, there is limited information on their relative utility. In this study, we compared the performance of different "classes" of genetic markers (mitochondrial DNA (mtDNA), nuclear DNA (nDNA), and allozymes) in terms of estimating levels and partitioning of genetic variation and of the relative accuracy and precision in estimating population allocations to mixed-stock fisheries. Individuals from eight populations of fall-run chum salmon ( Oncorhynchus keta ) from the Yukon River in Alaska and Canada were assayed at 25 loci. Significant differences in mitochondrial haplotype and nuclear allele frequencies were observed among five drainages. Populations from the U.S.-Canada border region were not clearly distinguishable based on multilocus allele frequencies. Although estimates of total genetic diversities were higher for the DNA loci ( H t = 0.592 and h = 0.647 for nDNA and mtDNA, respectively) compared with protein allozymes ( H t = 0.250), estimates of the extent of population differentiation were highly concordant across marker classes (mean theta = 0.010, 0.011, and 0.016 for allozymes, nDNA, and mtDNA, respectively). Simulations of mixed-stock fisheries composed of varying contributions of U.S. and Canadian populations revealed a consistent bias for overallocation of Canadian stocks when expected Canadian contributions varied from 0 to 40%, due primarily to misallocations among genetically similar border populations. No single marker class is superior for differentiating populations of this species at the spatial scale examined.

Yukon River

A method for nitrate collection for δ 15 N and δ 18 O analysis from waters with low nitrate concentrations

Recently, methods have been developed to analyze NO3- for &delta;15N and &delta;18O, improving our ability to identify NO3- sources and transformations. However, none of the existing methods are suited for waters with low NO3- concentrations (0.7-10 &micro;M). We describe an improved method for collecting and recovering NO3- on exchange columns. To overcome the lengthy collection loading times imposed by the large sample volumes (7-70 L), the sample was prefiltered (0.45 &micro;m) with a large surface area filter. Switching to AG2X anion resin and using a coarser mesh size (100-200) than previous methods also enhanced sample flow. Placement of a cation column in front of the anion column minimized clogging of the anion column by dissolved organic carbon (DOC) accumulation. This also served to minimize transfer of unwanted oxygen atoms from DOC to the 18O portion of the NO3- sample, thereby contaminating the sample and shifting &delta;18O. The cat-AG2X method is suited for on-site sample collection, making it possible to collect and recover NO3- from low ionic strength waters with modest DOC concentrations (80-800 &micro;M), relieves the investigator of transporting large volumes of water back to the laboratory, and offers a means of sampling rain, snow, snowmelt, and stream samples from access-limited sites.

Canadian Journal of Fisheries and Aquatic Sciences

Metal exposure in a benthic macroinvertebrate, Hydropsyche californica, related to mine drainage in the Sacramento River

A biomonitoring technique was employed to complement studies of metal transport in the upper Sacramento River affected by acid mine drainage. Metals (Al, Cd, Cu, Fe, Hg, Pb, and Zn) were determined in a resident invertebrate, Hydropsyche californica (Insecta: Trichoptera), and streambed sediments (<62 µm) to assess metal contamination within a 111-km section of the river downstream of the mining area. Metals in H. californica also were interpreted to be broadly indicative of metal exposure in fish. Total Hg was determined in the whole body of the insect, whereas Al, Cd, Cu, Fe, Pb, and Zn were additionally separated into operationally defined cytosolic (used as an indicator of exposure to bioavailable metal) and particulate fractions. Total concentrations of Cd, Cu, Hg, Pb, and Zn in sediments were consistent with documented upstream sources of acid mine drainage. Metal distribution patterns in H. californica and sediments were generally consistent for Cd, Cu, and Pb but inconsistent for Hg and Zn. Concentrations in H. californica indicated that bioavailable Cd, Cu, Pb, and Zn was transported at least 120 km downstream of the mine sources. Zinc in H. californica was elevated, but unlike sediments, did not decrease downstream. Mercury in H. californica was not elevated.

California

Direct behavioral evidence that unique bile acids released by larval sea lamprey (Petromyzon marinus) function as a migratory pheromone

Four behavioral experiments conducted in both the laboratory and the field provide evidence that adult sea lamprey ( Petromyzon marinus ) select spawning rivers based on the odor of larvae that they contain and that bile acids released by the larvae are part of this pheromonal odor. First, when tested in a recirculating maze, migratory adult lamprey spent more time in water scented with larvae. However, when fully mature, adults lost their responsiveness to larvae and preferred instead the odor of mature individuals. Second, when tested in a flowing stream, migratory adults swam upstream more actively when the water was scented with larvae. Third, when migratory adults were tested in a laboratory maze containing still water, they exhibited enhanced swimming activity in the presence of a 0.1 nM concentration of the two unique bile acids released by larvae and detected by adult lamprey. Fourth, when adults were exposed to this bile acid mixture within flowing waters, they actively swam into it. Taken together, these data suggest that adult lamprey use a bile acid based larval pheromone to help them locate spawning rivers and that responsiveness to this cue is influenced by current flow, maturity, and time of day. Although the precise identity and function of the larval pheromone remain to be fully elucidated, we believe that this cue will ultimately prove useful as an attractant in sea lamprey control.

Canadian Journal of Fisheries and Aquatic Sciences

Modeling data from double-tagging experiments to estimate heterogeneous rates of tag shedding in lake trout (Salvelinus namaycush)

Data from mark-recapture studies are used to estimate population rates such as exploitation, survival, and growth. Many of these applications assume negligible tag loss, so tag shedding can be a significant problem. Various tag shedding models have been developed for use with data from double-tagging experiments, including models to estimate constant instantaneous rates, time-dependent rates, and type I and II shedding rates. In this study, we used conditional (on recaptures) multinomial models implemented using the program SURVIV (G.C. White. 1983. J. Wildl. Manage. 47: 716-728) to estimate tag shedding rates of lake trout (Salvelinus namaycush) and explore various potential sources of variation in these rates. We applied the models to data from several long-term double-tagging experiments with Lake Superior lake trout and estimated shedding rates for anchor tags in hatchery-reared and wild fish and for various tag types applied in these experiments. Estimates of annual tag retention rates for lake trout were fairly high (80-90%), but we found evidence (among wild fish only) that retention rates may be significantly lower in the first year due to type I losses. Annual retention rates for some tag types varied between male and female fish, but there was no consistent pattern across years. Our estimates of annual tag retention rates will be used in future studies of survival rates for these fish.

Canadian Journal of Fisheries and Aquatic Sciences

Dynamics of the recovery of the western Lake Erie walleye (Stizostedion vitreum vitreum) stock

After its 1957 collapse under intensive fishing and environmental stresses, the walleye (Stizostedion vitreum vitreum) stock of western Lake Erie remained low throughout the 1960s. A moratorium on both sport and commercial fishing, resulting from the 1970 discovery of mercury concentrations in walleye flesh, provided an opportunity for the development of an international interagency management plan. The quota management plan developed depended on sequential projection of the fishable stock on the basis of estimated annual recruitment and reports of total withdrawals from the stock. The fishery reopened gradually and quota management (including allocation among jurisdictions) was implemented in 1976. The stock, which had been gradually increasing as a result of relatively strong year-classes produced in 1970, 1972, and 1974, responded well to limited exploitation and produced a record year-class in 1977. Quotas were exceeded in 1978-80, but the stock continued to improve to the extent that the recommended rate of exploitation was increased in 1980 and again in 1981. As the population expanded, growth began to decline; the decline became apparent in young-of-the-year in the early 1970s and in older walleyes in the late 1970s. This trend toward progressively slower growth, which continued in the 1977 and subsequent year-classes, was accompanied by an increase in length at sexual maturity and a decrease in the percentage of female walleyes reaching sexual maturity at age III. As a net result of these changes, the proportion of mature females in the stock (an index of stock fecundity) decreased slightly during the interval 1975-84, while the estimated biomass of the standing stock rose from 9 000 to nearly 26 000 t. Both sport and commercial catches increased markedly after 1980 in Lake Erie's central basin.

Canadian Journal of Fisheries and Aquatic Sciences

Changes in the nearshore and offshore zooplankton communities in Lake Ontario: 1981-88

We examined trends and factors influencing changes in nearshore and offshore zooplankton abundance and composition in Lake Ontario between 1981 and 1988. In the nearshore (southshore and eastern basin), zooplankton abundance decreased and shifts occurred in the relative abundances of Bosmina longirostris and Daphnia retrocurva (eastern basin) and Daphnia retrocurva and Daphnia galeata mendotae (southshore). These changes could have resulted from increased vertebrate predation or reduced food resources which intensified the effects of predation. In the offshore, the first appearance (FA) of the larger, less common cladoceran species occurred earlier in the season as of 1985. FA was correlated with cumulative epilimnetic temperature (CET) and the catch per unit effort (CPUE) of alewife (Alosa pseudoharengus) a?Y165 mm caught in U.S. waters in the spring. In 1987, when CET was high and CPUE of alewife a?Y165 mm was low, large populations of these cladocerans developed in June and July. Bythotrephes cederstroemi, a recent invader in the Great Lakes, was abundant only in 1987 when the CPUE of alewife was lowest. Changes in zooplankton abundance, development, and composition along the nearshore-offshore gradient reflected effects of temperature, habitat, and planktivory on the community.

Canadian Journal of Fisheries and Aquatic Sciences

Use of zooplankton to assess the movement and distribution of alewife (Alosa pseudoharengus) in south-central Lake Ontario in spring

Data from assessments of fish and zooplankton conducted during April and May-June 1986-88 in south-central Lake Ontario were examined for evidence that zooplankton size structure can be used to follow the movement of alewife (Alosa pseudoharengus). The spring influx of alewife into nearshore waters was linked with water temperature and coincided with a decline in the mean length of crustacean zooplankton and the virtual disappearance of zooplankters a?Y 0.9 mm. Alewife moving inshore to spawn fed heavily on the largest zooplankters, negating the possibility that changes in zooplankton size were wholly a response to seasonal recruitment as waters warm and the competition shifts to Bosmina. Offshore, there was usually no significant (P < 0.05) change in mean lengths of zooplankton in the upper water column between April and May-June, and zooplankters a?Y 0.9 mm always remained abundant, suggesting that few alewife were there from April through mid-June. We conclude that in large freshwater lakes where a planktivore is abundant, yet spatially concentrated, changes in size of crustacean zooplankton can facilitate understanding of the fish's movement and distribution.

Canadian Journal of Fisheries and Aquatic Sciences