Ron A. Heintz

Marine Subsidies in Freshwater: Effects of Salmon Carcasses on Lipid Class and Fatty Acid Composition of Juvenile Coho Salmon

Abstract Returning adult salmon represent an important source of energy, nutrients, and biochemicals to their natal streams and may therefore have a quantitative effect on the energy levels of stream-resident salmonids. We tested this hypothesis by constructing simulated streams for coho salmon Oncorhynchus kisutch to which we added 0, 1, and 4 carcasses/m2 (0, 0.71, and 2.85 kg wet mass/m2) of pink salmon O. gorbuscha. After 60 d we evaluated the lipid class and fatty acid composition of rearing coho salmon from the simulated streams; the lipid content and triacylglycerols of the coho salmon increased with increasing carcass density whereas phospholipids decreased. Increased amounts of triacylglycerols accounted for most of the lipid increase. In addition to increasing in concentration, the fatty acid composition of the triacylglycerols also changed with carcass density. Triacylglycerols of juvenile coho salmon from the control streams had significantly higher omega-3 : omega-6 ratios as a result of five...

MONITORING POLYNUCLEAR AROMATIC HYDROCARBONS IN AQUEOUS ENVIRONMENTS WITH PASSIVE LOW-DENSITY POLYETHYLENE MEMBRANE DEVICES

Low-density polyethylene membranes, typically filled with triolein, have been previously deployed as passive environmental samplers designed to accumulate nonpolar hydrophobic chemicals from water, sediments, and air. Hydrocarbons in such samplers, known as semipermeable membrane devices (SPMDs), diffuse through pores in the membranes and are trapped in the central hydrocarbon matrix, mimicking uptake by living organisms. Here, we describe laboratory and field verification that low-density polyethylene membrane devices (PEMDs) without triolein provide reliable, relatively inexpensive, time-integrated hydrocarbon sampling from water. For comparison, polynuclear aromatic hydrocarbon (PAH) uptake in SPMDs and pink salmon eggs also was studied. Total concentrations of PAH accumulated by PEMDs were highly correlated with concentrations in water (r2 > or = 0.99) and linear over the range tested (0-17 microg/L). Higher-molecular-mass PAH preferentially accumulated in PEMDs and in pink salmon eggs, but the source of oil in PEMDs remained identifiable. Accumulations of PAH were highly similar to those in SPMDs. The PEMDs retained approximately 78% of accumulated total PAH for 40 d in clean water. Thus, a simple plastic membrane can be conveniently used for environmental monitoring, particularly during situations in which contaminant concentrations are low (in the parts-per-billion range), variable, and intermittent.

Contaminant exposure and effects in pinnipeds: implications for Steller sea lion declines in Alaska

After nearly 3 decades of decline, the western stock of Steller sea lions (SSL; Eumetopias jubatus) was listed as an endangered species in 1997. While the cause of the decline in the 1970s and 1980s has been attributed to nutritional stress, recent declines are unexplained and may result from other factors including the presence of environmental contaminants. SSL tissues show accumulation of butyltins, mercury, PCBs, DDTs, chlordanes and hexachlorobenzene. SSL habitats and prey are contaminated with additional chemicals including mirex, endrin, dieldrin, hexachlorocyclohexanes, tetrachlorodibenzo-p-dioxin (TCDD) and related compounds, cadmium and lead. In addition, many SSL haulouts and rookeries are located near other hazards including radioactivity, solvents, ordnance and chemical weapon dumps. PCB and DDT concentrations measured in a few SSL during the 1980s were the highest recorded for any Alaskan pinniped. Some contaminant exposures in SSL appear to be elevated in the Gulf of Alaska and Bering Sea compared to southeast Alaska, but there are insufficient data to evaluate geospatial relationships with any certainty. Based on very limited blubber data, current levels of PCBs may not pose a risk to SSL based on comparison to immunotoxicity tissue benchmarks, but SSL may have been at risk from pre-1990 PCB exposures. While exposure to PCBs and DDTs may be declining, SSL are likely exposed to a multitude of other contaminants that have not been monitored. The impacts of these exposures on SSL remain unknown because causal effects have not been established. Field studies with SSL have been limited in scope and have not yet linked contaminant exposures to adverse animal health or population effects. Several biomarkers may prove useful for monitoring exposure and additional research is needed to evaluate their utility in SSL. We conclude that there are insufficient data to reject the hypothesis that contaminants play a role in the continued decline of SSL, and suggest that a coordinated monitoring program be developed which can be related to key biological, ecological and laboratory toxicity data.

Spatial Match-Mismatch between Juvenile Fish and Prey Provides a Mechanism for Recruitment Variability across Contrasting Climate Conditions in the Eastern Bering Sea

Understanding mechanisms behind variability in early life survival of marine fishes through modeling efforts can improve predictive capabilities for recruitment success under changing climate conditions. Walleye pollock (Theragra chalcogramma) support the largest single-species commercial fishery in the United States and represent an ecologically important component of the Bering Sea ecosystem. Variability in walleye pollock growth and survival is structured in part by climate-driven bottom-up control of zooplankton composition. We used two modeling approaches, informed by observations, to understand the roles of prey quality, prey composition, and water temperature on juvenile walleye pollock growth: (1) a bioenergetics model that included local predator and prey energy densities, and (2) an individual-based model that included a mechanistic feeding component dependent on larval development and behavior, local prey densities and size, and physical oceanographic conditions. Prey composition in late-summer shifted from predominantly smaller copepod species in the warmer 2005 season to larger species in the cooler 2010 season, reflecting differences in zooplankton composition between years. In 2010, the main prey of juvenile walleye pollock were more abundant, had greater biomass, and higher mean energy density, resulting in better growth conditions. Moreover, spatial patterns in prey composition and water temperature lead to areas of enhanced growth, or growth ‘hot spots’, for juvenile walleye pollock and survival may be enhanced when fish overlap with these areas. This study provides evidence that a spatial mismatch between juvenile walleye pollock and growth ‘hot spots’ in 2005 contributed to poor recruitment while a higher degree of overlap in 2010 resulted in improved recruitment. Our results indicate that climate-driven changes in prey quality and composition can impact growth of juvenile walleye pollock, potentially severely affecting recruitment variability.

Salmon Carcasses Increase Stream Productivity More than Inorganic Fertilizer Pellets: A Test on Multiple Trophic Levels in Streamside Experimental Channels

Abstract Inorganic nutrient amendments to streams are viewed as possible restoration strategies for re-establishing nutrients and stream productivity throughout the western coast of North America, where salmon runs and associated marine-derived nutrient subsidies have declined. In a mesocosm experiment, we examined the short-term (6 weeks) comparative effects of artificial nutrient pellets and salmon carcasses, alone (low and high amounts) and in combination, on stream food webs. Response variables included dissolved nutrient concentrations, biofilm ash-free dry mass (AFDM) and chlorophyll-a levels, macroinvertebrate density, growth and body condition of juvenile coho salmon Oncorhynchus kisutch, and whole-body lipid content of invertebrates and juvenile coho salmon. Most of the response variables were significantly influenced by carcass treatment; the only response variable significantly influenced by fertilizer pellet treatment was soluble reactive phosphorus (SRP) concentration. Ammonium-nitrogen conce...

Seasonal persistence of marine-derived nutrients in south-central Alaskan salmon streams

Spawning salmon deliver annual pulses of marine-derived nutrients (MDN) to riverine ecosystems around the Pacific Rim, leading to increased growth and condition in aquatic and riparian biota. The influence of pulsed resources may last for extended periods of time when recipient food webs have effective storage mechanisms, yet few studies have tracked the seasonal persistence of MDN. With this as our goal, we sampled stream water chemistry and selected stream and riparian biota spring through fall at 18 stations (in six watersheds) that vary widely in spawner abundance and at nine stations (in three watersheds) where salmon runs were blocked by waterfalls. We then developed regression models that related dissolved nutrient concentrations and biochemical measures of MDN assimilation to localized spawner density across these 27 stations. Stream water ammonium-N and orthophosphate-P concentrations increased with spawner density during the summer salmon runs, but responses did not persist into the following fall. The effect of spawner density on δ15N in generalist macroinvertebrates and three independent MDN metrics (δ15N, δ34S, and ω3:ω6 fatty acids) in juvenile Dolly Varden (Salvelinus malma) was positive and similar during each season, indicating that MDN levels in biota increased with spawner abundance and were maintained for at least nine months after inputs. Delta 15N in a riparian plant, horsetail (Equisetum fluviatile), and scraper macroinvertebrates did not vary with spawner density in any season, suggesting a lack of MDN assimilation by these lower trophic levels. Our results demonstrate the ready assimilation of MDN by generalist consumers and the persistence of this pulsed subsidy in these organisms through the winter and into the next growing season.

Straying of Adult Pink Salmon from their Natal Stream following Embryonic Exposure to Weathered Exxon Valdez Crude Oil

Abstract Numbers of strays (adult salmon returning to a nonnatal stream), straying rates, and distribution of strays were estimated for pink salmon incubated in oil-contaminated gravel and for an unexposed control group. The treatment groups were incubated in oiled gravel, which resulted in initial aqueous exposures for total polynuclear aromatic hydrocarbons (TPAHs) of 5 and 19 μg/L for a low and high dose, respectively; the control group was incubated in gravel without oil. Fry from each treatment group were marked with coded wire tags (CWTs). The numbers of tagged fish released were 65,409 from the control; 69,441 from the low-dose group; and 70,414 from the high-dose group. A total of 288,492 pink salmon were sampled for CWTs when returning to spawn in the natal stream of the experimental fish, when returning to streams within 60 km of the natal stream, and when returning to two hatcheries within 100 km of the natal stream. The frequencies of observed strays were 0.023, 0.030, and 0.025 for the contro...

Identification of Marine-Derived Lipids in Juvenile Coho Salmon and Aquatic Insects through Fatty Acid Analysis

Abstract The energetic benefits enjoyed by consumers in streams with salmon runs depend on how those benefits are accrued. Adult Pacific salmon Oncorhynchus spp. deliver significant amounts of nutrients (i.e., nitrogen and phosphorus) and carbon to streams when they spawn and die; these nutrient additions can have demonstrable effects on primary production in streams. Consumption of carcass tissues or eggs provides for direct energy subsidies to consumers and may have significant effects on their condition. In this study, comparisons of juvenile coho salmon O. kisutch and aquatic insects exposed to terrestrial and marine energy sources demonstrated that direct consumption of marine-derived lipids had a significant effect on the lipid reserves of consumers. Direct consumption of marine-derived tissues was verified through fatty acid analysis. Selected aquatic insects and juvenile coho salmon were reared for 6 weeks in experimental streams supplied with terrestrial or marine energy sources. Chironomid midge...

Energetic Cost of Ichthyophonus Infection in Juvenile Pacific Herring (Clupea pallasii)

The energetic costs of fasting and Ichthyophonus infection were measured in juvenile Pacific herring (Clupea pallasii) in a lab setting at three temperatures. Infected herring incurred significant energetic costs, the magnitude of which depended on fish condition at the time of infection (fat versus lean). Herring that were fed continually and were in relatively good condition at the time of infection (fat) never stored lipid despite ad libitum feeding. In feeding herring, the energetic cost of infection was a 30% reduction in total energy content relative to controls 52 days post infection. Following food deprivation (lean condition), infection caused an initial delay in the compensatory response of herring. Thirty-one days after re-feeding, the energetic cost of infection in previously-fasted fish was a 32% reduction in total energy content relative to controls. Body composition of infected herring subsequently recovered to some degree, though infected herring never attained the same energy content as their continuously fed counterparts. Fifty-two days after re-feeding, the energetic cost of infection in previously-fasted fish was a 6% reduction in total energy content relative to controls. The greatest impacts of infection occurred in colder temperatures, suggesting Ichthyophonus-induced reductions in body condition may have greater consequences in the northern extent of herrings range, where juveniles use most of their energy reserves to survive their first winter.

Return of warm conditions in the southeastern Bering Sea: Phytoplankton - Fish

In 2014, the Bering Sea shifted back to warmer ocean temperatures (+2 oC above average), bringing concern for the potential for a new warm stanza and broad biological and ecological cascading effects. In 2015 and 2016 dedicated surveys were executed to study the progression of ocean heating and ecosystem response. We describe ecosystem response to multiple, consecutive years of ocean warming and offer perspective on the broader impacts. Ecosystem changes observed include reduced spring phytoplankton biomass over the southeast Bering Sea shelf relative to the north, lower abundances of large-bodied crustacean zooplankton taxa, and degraded feeding and body condition of age-0 walleye pollock. This suggests poor ecosystem conditions for young pollock production and the risk of significant decline in the number of pollock available to the pollock fishery in 2–3 years. However, we also noted that high quality prey, large copepods and euphausiids, and lower temperatures in the north may have provided a refuge from poor conditions over the southern shelf, potentially buffering the impact of a sequential-year warm stanza on the Bering Sea pollock population. We offer the hypothesis that juvenile (age-0, age-1) pollock may buffer deleterious warm stanza effects by either utilizing high productivity waters associated with the strong, northerly Cold Pool, as a refuge from the warm, low production areas of the southern shelf, or by exploiting alternative prey over the southern shelf. We show that in 2015, the ocean waters influenced by spring sea ice (the Cold Pool) supported robust phytoplankton biomass (spring) comprised of centric diatom chains, a crustacean copepod community comprised of large-bodied taxa (spring, summer), and a large aggregation of midwater fishes, potentially young pollock. In this manner, the Cold Pool may have acted as a trophic refuge in that year. The few age-0 pollock occurring over the southeast shelf consumed high numbers of euphausiids which may have provided a high quality alternate prey. In 2016 a retracted Cold Pool precluded significant refuging in the north, though pollock foraging on available euphausiids over the southern shelf may have mitigated the effect of warm waters and reduced large availability of large copepods. This work presents the hypothesis that, in the short term, juvenile pollock can mitigate the drastic impacts of sustained warming. This short-term buffering, combined with recent observations (2017) of renewed sea ice presence over southeast Bering Sea shelf and a potential return to average or at least cooler ecosystem conditions, suggests that recent warm year stanza (2014–2016) effects to the pollock population and fishery may be mitigated.