Change in feeding ecology and trophic dynamics of Pacific salmon (Oncorhynchus spp.) in the central Gulf of Alaska in relation to climate events

The effects of climate events on the feeding ecology and trophic dynamics of Pacific salmon (Oncorhynchus spp.) in offshore waters of the central Gulf of Alaska were investigated during early summers (1994–2000), based on analyses of stomach contents, and carbon and nitrogen stable isotopes (δ¹³C and δ¹⁵N). Gonatid squids (mainly Berryteuthis anonychus) were the dominant prey of all salmon species except for chum salmon (O. keta). During the 1997 El Niño event and the 1999 La Niña event, squids decreased sharply in the diets of all Pacific salmon except coho salmon (O. kisutch) in the Subarctic Current, and chum salmon diets changed from gelatinous zooplankton (1995–97) to a more diverse array of zooplankton species. A δ¹³C and δ¹⁵N analysis indicated that all salmon species occupied the same branch of the food web in 1999–2000. We hypothesize that high‐seas salmon adapt to climate‐induced changes in their prey resources by switching their diets either within or between trophic levels. To understand the effects of climate change on Pacific salmon in the Gulf of Alaska, biological oceanographic research on B. anonychus and other important prey resources is needed.     

Growth, feed conversion and chemical composition of Atlantic salmon (Salmo salar L.) and Atlantic halibut (Hippoglossus hippoglossus L.) fed diets supplemented with krill or amphipods

The effects of partial replacement of fish meal (FM) with meal made from northern krill (Thysanoessa inermis), Antarctic krill (Euphausia superba) or Arctic amphipod (Themsto libellula) as protein source in the diets for Atlantic salmon (Salmo salar L.) and Atlantic halibut (Hippoglossus hippoglossus L.) on growth, feed conversion, macro‐nutrient utilization, muscle chemical composition and fish welfare were studied. Six experimental diets were prepared using a low‐temperature FM diet as control. The other diets included northern krill where 20, 40 or 60% of the dietary FM protein was replaced with protein from northern krill, and two diets where the FM protein was replaced with protein from Antarctic krill or Arctic amphipod at 40% protein replacement level. All diets were iso‐nitrogenous and iso‐caloric. Atlantic salmon grew from 410 g to approximately 1500 g during the 160 day experiment, and Atlantic halibut grew from 345 g to 500–600 g during the 150 day experiment. Inclusion of krill in the diets enhanced specific growth rate in salmon, especially during the first 100 days (P < 0.01), and in a dose–response manner in halibut for over the 150 day feeding period (P < 0.05). Feed conversion ratio did not differ between dietary treatments, and no difference was found in dry matter digestibility, protein digestibility and fish muscle composition. Good growth rates, blood parameters within normal ranges and low mortalities in all experimental treatments indicted that fish health was not affected either Atlantic salmon or Atlantic halibut fed the various zooplankton diets.     

Potential trophodynamic and environmental drivers of steelhead (Oncorhynchus mykiss) productivity in the North Pacific Ocean

Information on prey availability, diets, and trophic levels of fish predators and their prey provides a link between physical and biological changes in the ecosystem and subsequent productivity (growth and survival) of fish populations. In this study two long‐term data sets on summer diets of steelhead (Oncorhynchus mykiss) in international waters of the central North Pacific Ocean (CNP; 1991–2009) and Gulf of Alaska (GOA; 1993–2002) were evaluated to identify potential drivers of steelhead productivity in the North Pacific. Stable isotopes of steelhead muscle tissue were assessed to corroborate the results of stomach content analysis. We found the composition of steelhead diets varied by ocean age group, region, and year. In both the GOA and CNP, gonatid squid (Berryteuthis anonychus) were the most influential component of steelhead diets, leading to higher prey energy densities and stomach fullness. Stomach contents during an exceptionally warm year in the GOA and CNP (1997) were characterized by high diversity of prey with low energy density, few squid, and a large amount of potentially toxic debris (e.g., plastic). Indicators of good diets (high proportions of squid and high prey energy density) were negatively correlated with abundance of wild populations of eastern Kamchatka pink salmon (O. gorbuscha) in the CNP. In conclusion, interannual variations in climate, abundance of squid, and density‐dependent interactions with highly‐abundant stocks of pink salmon were identified as potential key drivers of steelhead productivity in these ecosystems. Additional research in genetic stock identification is needed to link these potential drivers of productivity to individual populations.     

Escaped farmed Atlantic salmon, Salmo salar L., feeding in Scottish coastal waters

Escaped reared salmon, Salmo salar L., were distinguished from wild salmon in the catch of a coastal salmon fishery on the west coast of Scotland. The stomach contents of 54 escaped fish were examined to determine their recent feeding history and 19 (35%) were found to contain food. The predominant prey were juvenile whiting, Merlangius merlangus (L.), unidentified Gadidae and sandeels (Ammodytidae), although other fish and invertebrates, mainly post-larval hermit crabs (Paguridae), were recorded. All these prey are pelagic or semi-pelagic. These observations demonstrate that escaped salmon feed on natural prey in coastal waters and extend our knowledge of the diet of salmon in their marine phase.     

Using stable isotope data to quantify niche overlap and diets of muskellunge,northern pike and walleye in a deep Minnesota lake

Muskellunge (Esox masquinongy Mitchill), northern pike (Esox lucius L.) and walleye (Sander vitreus Mitchill) often coexist in lake communities, yet uncertainty exists about the potential for interspecific competition among these top predators. Stable isotope data were used to assess niche overlap and diets of these predators in Elk Lake (Minnesota, U.S.A). δ¹³C indicated primary production sources (e.g. pelagic v. littoral) and δ¹⁵N indicated trophic position; the bivariate distribution of these isotopes defined the species’ isotopic niche. Niche overlap probabilities were calculated and stable isotope mixing models were used to quantify diet proportions. Muskellunge and northern pike niches overlapped little (<10%), while walleye overlapped muskellunge (15%–60%) and northern pike (33%–53%) more extensively. Muskellunge diets focused (50%) on cisco (Coregonus artedi Lesueur), walleye primarily assimilated non-cisco prey fish (80%), and northern pike diets were dominated by non-cisco prey fish (45%) and invertebrates (40%). The presence of a cisco population and the flexibility of northern pike to use invertebrate resources may decrease potential competition among these predators. However, cisco are threatened by climate change and eutrophication, and our results suggest that extirpation of cisco may cause major changes in potential competitive interactions among these top predators. Moreover, cisco were unique among prey species in their ability to exploit pelagic energy, such that loss of cisco will likely alter energy flow in lake food webs where they currently exist.     

The effect of dietary krill supplementation on epithelium-associated bacteria in the hindgut of Atlantic salmon (Salmo salar L.): a microbial and electron microscopical study

Atlantic salmon (Salmo salar L.) were fed fishmeal protein for 46 days, and 500 g kg⁻¹ of fishmeal protein substituted with meal from Northern krill (Meganyctiphanes norvegica). No differences were observed in weight gain, length gain, feed conversion or specific growth rate between the groups that could be attributed to dietary manipulation. The adherent microbiota in the hindgut of the two rearing groups were further investigated. By substituting fishmeal with krillmeal, the total viable counts of aerobic and facultative aerobic bacteria colonizing the hindgut of Atlantic salmon increased from 8.5 × 10⁴ to 2.2 × 10⁶. Furthermore, dietary krillmeal affected the adherent hindgut microbiota. The Gram‐positive bacteria Carnobacteria piscicola, Microbacterium oxydans, Microbacterium luteolum and Staphylococcus equorum spp. linens and the Gram‐negatives Psychrobacter spp. and Psychrobacter glacincola were not isolated from hindgut of fish fed the krill diet. On the other hand, Pseudomonas fulgida, Pseudomonas reactans and Stenotrophomonas maltophila were not isolated from the control group fed fishmeal. Acinetobacter lwoffi, which is not normally found in the fish gut, was isolated from both feeding groups. Transmission electron microscopy showed bacteria‐like profiles between the hindgut microvilli in both feeding groups indicating autochthonous microbiota. When fish were fed the krill diet, hindgut enterocytes were replete with numerous irregular vacuoles. These vacuoles were not observed in fish fed the fishmeal protein.     

Juvenile Chinook salmon,Oncorhynchus tshawytscha,growth and diet in riverine habitat engineered to improve conditions for spawning

Abstract Many habitat enhancement techniques aimed at restoring salmonid populations have not been comprehensively assessed. The growth and diet of juvenile Chinook salmon, Oncorhynchus tshawytscha (Walbaum), rearing in a reach designed to enhance spawning were evaluated to determine how a non‐target life stage fared in the engineered habitat. Prior work demonstrated differences in food web structure between restored and unenhanced reaches of the Merced River, thus juvenile salmon feeding dynamics were also hypothesised to vary. Dependent variables were compared among fish collected from within and near the upper boundary of the restored reach and in an unenhanced habitat upstream. Diets, otolith‐derived growth and stable isotope‐inferred trophic positions were compared. Baetidae mayflies were particularly important prey in the restored reach, while elsewhere individuals exhibited heterogeneous diets. Salmon residing at the bottom of the restored reach exhibited slightly faster growth rates relative to fish collected elsewhere, although stable isotope and diet analyses suggested that they fed at a relatively low trophic position. Specialised Baetis predation and/or abundant interstitial refugia potentially improved rearing conditions in the restored reach. Data suggest that gravel enhancement and channel realignment designed to augment adult spawning habitat may simultaneously support juvenile Chinook salmon despite low invertebrate food resources.     

Fillet Quality and Processing Attributes of Postsmolt Atlantic Salmon,Salmo salar,Fed a Fishmeal‐free Diet and a Fishmeal‐based Diet in Recirculation Aquaculture Systems

Many studies have evaluated the adequacy of alternate ingredient diets for Atlantic salmon, Salmo salar, mainly with focus on fish performance and health; however, comprehensive analysis of fillet quality is lacking, particularly for salmon fed these diets in recirculation aquaculture systems (RAS). To this end, a study was conducted comparing fillet quality and processing attributes of postsmolt Atlantic salmon fed a fishmeal‐free diet (FMF) versus a standard fishmeal‐based diet, in replicate RAS. Mean weight of Atlantic salmon fed both diets was 1.72 kg following the 6‐mo trial and survival was >99%. Diet did not affect (P > 0.05) processing and fillet yields, whole‐body proximate composition(fat, moisture, protein), fillet proximate composition, cook yield, fillet texture, color, or omega‐3 fatty acid fillet content, including eicosapentaenoic acid and docosahexaenoic acid levels. Whole‐body ash content was greater in salmon fed the FMF diet. The FMF diet resulted in a wild fish‐in to farmed fish‐out ratio of 0:1 per Monterey Bay Aquarium's Seafood Watch criteria due to its fishmeal‐free status and use of lipids from fishery byproduct. Overall, fillet quality and processing attributes were generally unaffected when feeding a diet devoid of fishmeal to postsmolt Atlantic salmon cultured in RAS. [Correction added on 7 September 2017, after first online publication: the P value in Abstract has been changed from “P < 0.05” to “P > 0.05”.].     

The study of feeding preferences using radiography: oxytetracycline as a feeding deterrent and squid extract as a stimulant in diets for Atlantic salmon

A method for identifying and confirming the efficacy of feeding stimulants and deterrents is described. Feeding studies carried out using Atlantic salmon, Salmo salar L., have shown that the addition of squid extract as a stimulant to feeds may lead to modification of feed intake, and may ameliorate the negative effects of unpalatability when fish are presented with medicated diets.     

Effects of diet supplementation of soya‐saponins,isoflavones and phytosterols on Atlantic salmon (Salmo salar,L) fry fed from start‐feeding

A 14‐week trial was conducted to investigate the effects of antinutritional factors (ANFs) commonly present in soybean ingredients, singly and in combination, on Atlantic salmon (Salmo salar L.) fed from start‐feeding. The experimental diets consisted of a negative control fish meal diet (FM), and a positive control diet with 167 g kg^?1 soybean meal inclusion (SBM) and four diets based on the FM diet supplemented with 2 g kg^?1 soya‐saponins (SAP), 1.5 g kg^?1 isoflavones (IFL), 0.3 g kg^?1 phytosterols (PHS) or a mixture of these (MIX). Fish fed the SAP diet showed significantly higher growth performance than those fed FM, while the IFL treatment significantly decreased growth performance of salmon fry. Fish fed the IFL diet had significantly lower maltase activity and higher trypsin activity in proximal intestine than fish fed the FM diet. Histological differences were observed in the liver of fish fed the IFL diet, characterized by reduced size of the hepatocytes. Fish fed the PHS and IFL diets showed the highest frequencies of skeletal deformities among the six treatments. In conclusion, the results indicate that purified isoflavones may negatively affect growth performance, intestinal function, liver metabolism and bone formation of salmon fry.     

Intensive fishing has not forced dietary change in the South American fur seal Arctophoca (=Arctocephalus) australis off Río de la Plata and adjoining areas

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Effects of dietary protein, and fat level and rapeseed oil on growth and tissue fatty acid composition and metabolism in Atlantic salmon (Salmo salar L.) reared at low water temperatures

A 12‐week feeding trial was conducted to elucidate the interactive effects of dietary fat, protein contents and oil source on growth, whole body proximate composition, protein productive value (PPV) and fatty acid (FA) composition of muscle and liver in Atlantic salmon (Salmo salar L.)` at low water temperatures (4.2 °C). Triplicate groups of Atlantic salmon (initial weight 1168 g) were fed six isoenergetic diets, formulated to provide either 390 g kg⁻¹ protein and 320 g kg⁻¹ fat (high‐protein diets) or 340 g kg⁻¹ protein and 360 g kg⁻¹ fat (low‐protein diets). Within each dietary protein/fat level, crude rapeseed oil (RO) comprised 0, 30 or 60% (R0, R30, R60, respectively) of the added oil. After 12 weeks, the overall growth and feed conversion ratio (FCR) were very good for all treatments [thermal growth coefficient (TGC): 4.76 (±0.23); FCR: 0.85 (±0.02)]. Significant effects were shown owing to the oil source on specific growth rate and TGC only. The liver and muscle FA compositions were highly affected by the graded inclusion of RO. The PPV was significantly affected by the dietary protein level. The results of this study suggest that more sustainable, lower protein diets with moderate RO inclusion can be used in Atlantic salmon culture at low water temperatures with no negative effects on growth and feed conversion, no major detrimental effects on lipid and FA metabolism and a positive effect on protein sparing.     

Fatty acid signatures and stomach contents of four sympatric Lake Trout: assessment of trophic patterns among morphotypes in Great Bear Lake

Sympatric diversification in Lake Trout is generally linked to differences in habitat use (especially depth) as a result of foraging on different prey items. However, extensive sympatric divergence has taken place in the shallow waters (≤30 m) of Great Bear Lake, with multiple Lake Trout morphs varying in head and fin characteristics. To investigate diet partitioning as a potential explanatory mechanism for this diversification, we assessed trophic characteristics and relationships among four sympatric shallow‐water morphs of Lake Trout via analyses of fatty acids and stomach contents. Fatty acids and stomach contents both identified Lake Trout, Cisco and Mysis as key prey items in Lake Trout diets. Interestingly, terrestrial invertebrates were also seasonally important among morphs, reflecting temporal variability of available prey in this arctic lake. Some diet partitioning was observed among morphs; Morph 1 was characterised as a generalist, Morph 3 was more benthic‐oriented, and Morphs 2 and 4 were mainly pelagic feeders. Of the latter, Morph 4 was the most specialised, whereas Morph 2 exhibited alternative feeding tactics of benthic cannibalistic and pelagic piscivorous feeding. Our findings demonstrate that complementary dietary methods can elucidate habits of opportunistic feeders, a task that can often be problematic, given their complex and variable diets. Our results add new information and perspectives on the current model of Lake Trout differentiation, demonstrating niche partitioning based on benthic versus pelagic habitat use and generalist versus specialist feeding tactics.     

Foraging behaviour of juvenile pink salmon (Oncorhynchus gorbuscha) and size‐dependent predation risk

Two hypotheses related to effects of juvenile pink salmon (Oncorhynchus gorbuscha) foraging behaviour and size on their predation risk were evaluated using field data collected in Prince William Sound, Alaska 1 995–97. My results supported the hypothesis that low macrozooplankton density leads to dispersion of juvenile salmon from shallow nearshore habitats and greater predation risk, but zooplankton type was an important factor. When the biomass of large copepods (primarily Neocalanus spp.) declined, salmon dispersed from shallow nearshore habitats, and mean daily individual predator consumption of salmon increased by a factor of 5. A concomitant five‐fold increase in the probability of occurrence of salmon in predator stomachs supported the notion that increased predation on salmon was caused by a greater overlap between predator and prey when salmon dispersed offshore, not an increase in the number of salmon consumed per feeding bout. The results also generally supported the hypothesis that the timing of predation events modifies the nature of size‐dependent predation losses of salmon to different predator groups (small and large planktivores and piscivores). Size‐dependent vulnerabilities of salmon to predators were a function of both predator and prey sizes. When simulated predation was shifted from May to June, the vulnerability of salmon became more dependent on their growth than initial size. But, the size‐ and growth‐dependent vulnerabilities of salmon differed more among predator groups than between May and June, suggesting that changes in the composition of predator fields could more strongly affect the nature of size‐dependent predation than changes in the timing of predation losses.     

Foraging mode variation in three stream‐dwelling salmonid fishes

Despite long‐standing interest in foraging modes as an important element of animal space use, few studies document and compare individual foraging mode differences among species and ecological conditions in the wild. We observed and compared foraging modes of 61 wild Arctic charr, Salvelinus alpinus, 42 brown trout, Salmo trutta, and 50 Atlantic salmon, Salmo salar, in their first growing season over a range of habitats in 10 Icelandic streams. We found that although stream salmonids typically sit‐and‐wait to ambush prey from short distances, Arctic charr were more mobile during prey search and prior to prey attack than Atlantic salmon, whereas brown trout were intermediate. In all three species, individuals that were mobile during search were more likely to be moving when initiating attacks on prey, although the strength and the slope of this relationship differed among species. Arctic charr also differed from salmon and trout as more mobile individuals travelled longer distances during prey pursuits. Finally, coupled with published data from the literature, salmonid foraging mobility (both during search and prior to attack) clearly decreased from still water habitats (e.g., brook charr), to slow‐running waters (e.g., Arctic charr) to fast‐running waters (e.g., Atlantic salmon). Hence, our study suggests that foraging mode of young salmonids can vary distinctly among related species and furthers our understanding of the behavioural mechanisms shaping the geographical distribution of wild salmonids.     

Atlantic salmon, Salmo salar L., feed in Scottish coastal waters

Abstract. The stomach contents of 256 Atlantic salmon, Salmo salar L., of fork lengths 53–66cm, caught in bag nets on the Scottish west coast near Ullapool, were examined between June 1983 and July 1986. A total of 61 fish contained fish in the stomach or had faecal pellets containing fish bones in the gut. All recognizable whole fish were sandeels, Ammodytes marinus Raitt, ranging in size from 4.5 to 15cm. No evidence for crustacean or other non-fish prey items was found. Results indicate that feeding salmon were caught up to a certain cut-off point in June or early July, after which all salmon sampled were not feeding. It is suggested that either there is a local feeding stock of salmon or that fish feed during migration from the Faroe Isles or other possible distant water origins.     

Effects of feed processing conditions and acid salts on nutrient digestibility and physical quality of soy‐based diets for Atlantic salmon (Salmo salar)

Two experiments were conducted to investigate effects of feed processing conditions and potassium diformate (KDF) supplementation on apparent digestibility of nutrients in Atlantic salmon (Salmo salar) and physical quality of extruded feed. In Exp. 1, diets with raw or expander pretreated full‐fat soybean meal (FFSBM) at 100 or 120 °C were extruded at 110 °C. Expander pretreatment significantly (P < 0.05) improved the digestibility of arginine, glutamine and tyrosine in Atlantic salmon. The higher digestibility of expander pretreated FFSBM was confirmed in mink (Mustela vison). In Exp. 2, diets with defatted soybean meal (SBM) were extruded at 110, 130 or 150 °C. The results showed that increasing extrusion temperatures significantly (P < 0.05) improved the digestibility of most major nutrients and amino acids in Atlantic salmon. In general, KDF supplementation to FFSBM and SBM diets did not affect digestibility of nutrients in Atlantic salmon or mink. Expander pretreatment and increasing extrusion temperatures increased pellet expansion, while KDF supplementation reduced pellet expansion.     

Astaxanthin deposition in the flesh of Atlantic Salmon, Salmo salar L., in relation to dietary astaxanthin concentration and feeding period

Atlantic salmon, Salmo salar L., were fed nine experimental diets containing from 0 to 200 mg astaxanthin per kg^?1 for six time periods, ranging from 3 to 21 months, in sea cages at Matre Aquaculture Research Station, Matredal, Norway. The sampled fish had an initial mean weight of 115 g and reached a weight of 3.2 kg at the termination of the experiment. Every third month, 10 fish from each dose and time group were sampled and the astaxanthin concentration in the flesh determined. The amount of astaxanthin in the flesh ranged from 0.7 to 8.9 mg kg^?1 at the termination of the experiment. This paper discusses deposition of astaxanthin in the flesh of Atlantic salmon in relation to dietary carotenoid levels in the 0–200 mg kg^?1 range and feeding times of 3–21 months. Under the conditions of this experiment, no significant effect on astaxanthin deposition rate could be achieved by increasing the astaxanthin level above 60 mg kg dry feed^?1. Atlantic salmon should be fed astaxanthin-supplemented diets during the whole seawater stage in order to obtain maximal astaxanthin level in the flesh.     

Dietary menadione nicotinamide bisulphite (vitamin K3) does not affect growth or bone health in first-feeding fry of Atlantic salmon (Salmo salar L.)

The aim of the present study was to elucidate if increasing levels of added vitamin K in the feed influenced fish growth, health or the incidences of bone deformities in Atlantic salmon ( Salmo salar L.) from onset of start feeding to ∼100 g in freshwater. Duplicate groups of Atlantic salmon fry (∼0.20 g) were fed eight levels (0, 2.5, 5, 7.5, 10, 15, 20 and 50 mg menadione kg⁻¹) of the vitamin K derivate menadione nicotinamide bisulphite (MNB) in a regression design for 28 weeks. All fish maintained high growth rates throughout the experiment, and showed no significant differences in specific growth rate, condition factor, whole body proximate analysis, blood coagulation time, vertebra morphology or mechanical properties of vertebrae. We found a dose-response between whole body vitamin K concentration and the dietary MNB supplementation level. Analysis of liver γ-glutamylcarboxylase activity revealed significant dose-dependent differences between groups given the 0, 10 and 50 mg MNB kg⁻¹ diets. In conclusion, Atlantic salmon seems to require low levels of dietary vitamin K, and the amount of vitamin K found naturally in the presently used feed ingredients may be enough to maintain optimal growth, health and bone strength in Atlantic salmon fry from start feeding.     

Wild origin and enriched environment promote foraging rate and learning to forage on natural prey of captive reared Atlantic salmon parr

Abstract – After release to the wild, captive reared salmon have shown lower foraging rates on natural prey and impaired ability to avoid natural predators and thus lower survival compared with wild‐born conspecifics. Here, we examine whether captive breeding influences learning of foraging on natural prey and how enriched rearing methods may improve foraging on natural prey by Atlantic salmon (Salmo salar) parr. We reared offspring of hatchery or wild salmon of the same population in either a standard or enriched environment at production‐scale densities. The enriched environment featured submerged overhead shelter, varying water current, depth and direction and consequently alterations in food dispersion. Parr reared in the enriched environment expressed higher feeding rates, and parr of wild origin started to forage earlier on natural prey. The enriched method promoted foraging of hatchery reared parr on natural prey and is easily applicable to commercial production of salmonids for reintroduction or stock enhancement purposes.