Climate change-informed dietary modeling in Pacific cod: Experimentally-derived effects of temperature and dietary quality on carbon and nitrogen stable isotope trophic discrimination factors.

Stable isotope analysis is a powerful tool for dietary modeling and trophic ecology research. A crucial piece of information for isotopic dietary modeling is the accurate estimation of trophic discrimination factors (TDFs), or the isotopic offset between a consumer's tissue and its diet. In order to parameterize stable isotope dietary models for future climate scenarios, we investigated the effect of water temperature and dietary protein and lipid content on TDFs in juvenile Pacific cod (Gadus macrocephalus). Pacific cod are a commercially and ecologically important species, with stock numbers in the northeast Pacific recently having dropped by more than 70%. We tested four water temperatures (6, 8, 10, and 12°C) and two dietary regimens (low and high lipid content), representing a range of potential ocean temperature and prey quality scenarios, in order to determine carbon and nitrogen TDFs in juvenile Pacific cod. Additionally, we assessed dietary intake and proximate composition of the experimental fish in order to estimate consumption, assimilation, and retention of dietary nutrients. The results of this study suggest that dietary protein catabolism is a primary driver of nitrogen TDF variability in juvenile Pacific cod. Across all temperature treatments from 6 to 12°C, fish reared on the lower quality, lower lipid content diet had higher nitrogen TDFs. The mean TDFs for fish raised on the higher lipid, lower protein diet were +3.40 ‰ for nitrogen (Δ15N) and +0.36 ‰ for lipid-corrected carbon (Δ LC 13C). The mean TDFs for fish raised on the lower lipid, higher protein diet were +4.09 ‰ for nitrogen (Δ15N) and 0.00 ‰ for lipid-corrected carbon (Δ LC 13C). Lipid-corrected carbon isotope data showed that, regardless of temperature, fish consuming the lower lipid diet had essentially no trophic discrimination between diet and bulk tissues. We found no ecologically meaningful differences in TDFs due to water temperature across the 6°experimental range. The results of this experiment demonstrate that dietary quality, and more specifically the use of dietary protein for energetic needs, is a primary driver of trophic discrimination factors. The TDFs determined in this study can be applied to understanding trophic ecology in Pacific cod and closely related species under rapidly changing prey availability and ocean temperature conditions.

Effects of Environmental Temperature on the Dynamics of Ichthyophoniasis in Juvenile Pacific Herring (Clupea pallasii).

The effects of temperature and infection by Ichthyophonus were examined in juvenile Pacific herring (Clupea pallasii) maintained under simulated overwinter fasting conditions. In addition to defining parameters for a herring bioenergetics model (discussed in Vollenweider et al. this issue), these experiments provided new insights into factors influencing the infectivity and virulence of the parasite Ichthyophonus. In groups of fish with established disease, temperature variation had little effect on disease outcome. Ichthyophonus mortality outpaced that resulting from starvation alone. In newly infected fish, temperature variation significantly changed the mortality patterns related to disease. Both elevated and lowered temperatures suppressed disease-related mortality relative to ambient treatments. When parasite exposure dose decreased, an inverse relationship between infection prevalence and temperature was detected. These findings suggest interplay between temperature optima for parasite growth and host immune function and have implications for our understanding of how Ichthyophonus infections are established in wild fish populations.

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 herring's range, where juveniles use most of their energy reserves to survive their first winter.

Seasonal cycles in whole-body proximate composition and energy content of forage fish vary with water depth.

Quantifying the nutritional quality of forage fish is integral for understanding upper trophic levels as forage fish are the dominant prey for top predator fish, marine mammals, and sea birds. Many existing reports documenting body composition of forage species are not comparable due to confounding effects. This study systematically assessed the variability in proximate composition and energy content of 16 forage species in southeastern Alaska (57.2626 N/133.7394 W) between 2001 and 2004. Variation in energy and lipid contents was related to habitat, epipelagic planktivores varying most, mesopelagics intermediate, and demersal species relatively invariable. Season was the greatest source of variation as a result of short growing seasons at high latitude and energy allocation strategies for reproduction and growth. Among species that varied seasonally, energy and lipid increased over summer and declined during winter. Annual differences in body composition occurred during periods of peak energy content. Sampling recommendations and guidance for bioenergetics models are provided.

Changes in the lipid class and proximate compositions of coho salmon (Oncorhynchus kisutch) smolts infected with the nematode parasite Philonema agubernaculum.

Coho salmon (Oncorhynchus kisutch) smolts infected with the nematode Philonema agubernaculum had 36% lower mean lipid content (1.4%) than nonparasitized coho salmon (2.2%) harvested simultaneously from the same outmigration. Lengths, weights, and condition factors, as well as protein and moisture content, did not differ significantly between the two groups. Lipid class compositions differed significantly between the parasitized and nonparasitized fish. None of the nematode-infected fish contained detectable triacylglycerols (TAG) or monoacylglycerols (MAG). In contrast, mean TAG and MAG contents of the nonparasitized fish totaled 5.5% of the extracted lipid. Infected smolts had lower cholesterol contents than did uninfected coho (17% for infected, 33% for uninfected). Parasitized fish had significantly higher levels of free fatty acids (mean of 57% for parasitized vs. 35% for nonparasitized) as well as the phospholipids phosphatidylethanolamine (PE) and phosphatidylcholine (PC). However, the PC/PE ratios for infected and noninfected coho did not differ significantly (2.2 for infected vs. 2.0 for uninfected). These differences suggest that the parasitic nematodes are either harvesting storage energy directly from the coho or are placing additional energetic demands on the fish to cope with the infection.