Control of Salmincola californiensis (Copepoda: Lernaeapodidae) in rainbow trout, Oncorhynchus mykiss (Walbaum): a clinical and histopathological study

A stock of rainbow trout, Oncorhynchus mykiss, held at an experimental facility, was found to be heavily infested with the lernmaeapodid copepod Salmincola californiensis. The efficacy and effects of treatment were compared with ivermectin or manual removal of parasites as a means of control of S. californiensis. One group of fish was orally intubated with 0.2 mg ivermectin active ingredient kg-1 fish. A second treatment was administered after a further 14 days. In a second group of fish, parasites were manually removed from the gills using forceps. These fish were sampled for up to 21 days post-first removal of parasites. In the ivermectin-treated fish adult parasites became inactive and changed colour within 18 h of the initial treatment. Copepods began to disappear by day 3 post-treatment and by day 31 almost all embedded female parasites had disappeared. Gills were clinically normal apart from cavitation deformity resulting from parasite attachment. Post-ivermectin treatment, there was an increase in the number of eosinophilic granular cells surrounding the bulla of attached S. californiensis, but from day 31 post-treatment these were replaced by macrophages and epithelioid cells to form a necrotic focus. In manually picked fish there was extensive haemorrhage in the interlamellar spaces as a result of parasite removal. At sites of parasite removal tissue necrosis was minimal and healing was rapid. At the end of the sampling period the structure of the gill was improved. The use of oral dosage with ivermectin is an effective treatment for S. californiensis and could be particularly beneficial for use with endangered salmon broodstocks infested with the parasite.     

Importance of river migration to the development of seawater tolerance in Columbia River anadromous salmonids

Gradually increasing levels of gill Na⁺K⁺ ATPase activity were observed in juvenile chinook, Oncorhynchus tshawytscha, and coho, Oncorhynchus kisutch, salmon and steelhead trout, Salmo gairdneri, undergoing parr-smolt transformation in artificial rearing facilities on the Columbia River. Portions of the same populations released to migrate seaward, however, generally showed much greater increases in enzyme activity with time and distance from the release point. After migrating 714 km to the Columbia River estuary, spring chinook salmon had a mean gill Na⁺K⁺ ATPase activity 2.5 times greater than fish retained at the hatchery and 1.9 times greater than fish adapted to 28 ppt seawater for 208 days. Similar observations were made on coho salmon.     

Chicken-derived IgY recognizes developing and mature stages of Loma salmonae (Microsporidia) in Pacific salmon, Oncorhynchus spp.

A polyclonal IgY preparation, purified from yolk of immunized hens, is shown to be useful for the detection of Loma salmonae in Pacific salmon. Purity of the IgY was confirmed by SDS-PAGE and western blots of yolk protein extracts. Reactivity of the IgY preparation to L. salmonae was determined through indirect-fluorescent antibody tests on purified spores and by immunohistochemistry. Developing xenomas in histological sections of heart and gill tissue of infected chinook salmon (Oncorhynchus tshawytscha) were visualized as early as 4 weeks post-exposure (PE). Non-lethal detection of L. salmonae was obtained by indirect-fluorescent microscopy of acetone-fixed gill smears. The IgY preparation was shown to react with another Loma sp., L. branchialis. In contrast, no reactivity was observed with other microsporidian parasites of fish such as Pseudoloma neurophilia and Glugea anomala. Yolk derived IgY may be useful in the detection of other fish pathogens.     

Prey selectivity and diel feeding chronology of juvenile chinook (Oncorhynchus tshawytscha) and coho (O. kisutch) salmon in the Columbia River plume

We studied salmon feeding selectivity and diel feeding chronology in the Columbia River plume. Juvenile chinook and coho salmon were caught by trawling at 2–3 h intervals throughout a diel period on three consecutive days (21–23 June 2000) at stations located 14.8 and 37 km offshore from the mouth of the Columbia River. A total of 170 chinook salmon were caught at the inshore and 79 chinook and 98 coho salmon were caught at the offshore station. After each trawl, potential prey were sampled at different depths with 2–3 different types of nets (1‐m diameter ring net, bongo net, neuston net). Despite the variability in zooplankton abundance, feeding selectivity was surprisingly constant. Both salmon species fed selectively on larger and pigmented prey such as hyperiid amphipods, larval and juvenile fish, various crab megalopae, and euphausiids. Hyperiid amphipods were abundant in the salmon diets and we hypothesize that aggregations of gelatinous zooplankton may facilitate the capture of commensal hyperiid amphipods. Small copepods and calyptopis and furcilia stages of euphausiids dominated the prey field by numbers, but were virtually absent from salmon diet. Juvenile chinook salmon, with increasing body size, consumed a larger proportion of fish. Stomach fullness peaked during morning hours and reached a minimum at night, suggesting a predominantly diurnal feeding pattern. In general, both chinook and coho salmon appear to be selective, diurnal predators, preying mostly on large and heavily pigmented prey items, in a manner consistent with visually oriented, size‐selective predation.     

Distribution,abundance and growth of juvenile salmonids off the coast of Oregon and Washington,summer (1980)

Juvenile salmonids (< 50 cm) were sampled by purse seine off the Pacific coast from Tillamook Bay, Oregon, to Copalis Head, Washington, during the period May through September (1980). Temporal distribution and abundance of the major Columbia River species were determined. Spring chinook salmon (Oncorhynchus tshatuytscha) and steel head (Salmo gairdneri) were present only during early cruises and were distributed almost entirely in the Columbia River plume and the sample area to the north. Coho salmon (O. kisutch) and fall chinook salmon were distributed more uniformly throughout the sampling area and were relatively abundant throughout the sampling period. Concentrations of fish were found only within 28 km of the shore. A number of fish that had been marked before or during their outmigration from the Columbia River system were recaptured. It appeared possible that with concentrated sampling in areas with high fish abundances, sufficient numbers of marked juvenile salmonids could be captured to provide relative survival estimates between different stocks of Columbia River fish.     

The suitability of anadromous Arctic charr as host and vector of the monogenean Gyrodactylus salaris

Abstract –  In 2000, the Atlantic salmon ( Salmo salar L.) stock in the river Signaldalselva, North Norway was found to be infected with the monogenean Gyrodactylus salaris (Malmberg). This river system has a multispecies fish community of nine species in total. Low densities of Atlantic salmon parr were found during electrofishing in October (2003) with a prevalence of 94% and a mean abundance of 848 G. salaris. This watercourse also holds a riverine stock of anadromous Arctic charr ( Salvelinus alpinus ), which is not common in Norway. Among the parr of Arctic charr, 64% were infested with an abundance of 23 parasites per fish. The most heavily infected 0+ and 1+ juvenile Arctic charr had 469 and 534 parasites, respectively, indicating that the parasites are able to reproduce on Arctic charr in the wild. There was little variation in the rates of infestation of Arctic charr along the studied stretch of the river, even in areas with very low densities of Atlantic salmon parr. The parr of anadromous Arctic charr seem to be suitable as long-term hosts for G. salaris . Moreover, a few (22%) infested adult sea-running Arctic charr were captured after ascending the river in the autumn, which shows that large fish may also act as carriers of G. salaris . However, no parasites were recorded on potential anadromous Arctic charr before descending during early spring (April 2003 and April 2004). Thus, it is still not clear whether anadromous Arctic charr are able to disperse the parasite between watercourses. So far, it is uncertain whether G. salaris may increase the mortality rates of Arctic charr and thereby be a threat to these unusual occurring riverine anadromous Arctic charr stocks.     

The effects of the 1983 El Niño on Oregon's Coho (Oncorhynchus kisutch) and Chinook (O. tshawytscha) Salmon

The 1983 El Niño event off the Pacific Coast of North America resulted in increased adult mortality and decreased average size for Oregon's coho and chinook salmon. Actual return of adult coho salmon to the Oregon Production Area in 1983 was only 42% of the pre-season prediction. Coho smolts entering the ocean in the spring of 1983 also survived poorly, resulting in low adult returns again in 1984. Abundance of chinook stocks in southern Oregon was also reduced, as was abundance of Columbia River chinook stocks that show localized ocean distribution. Northerly migrating chinook stocks from the Columbia River showed little or no decline in abundance. The average weight of coho and chinook salmon landed in 1983 by Oregon's commercial troll fishery was the lowest recorded since statistics were first recorded in 1952. Comparison of the length-weight relationship for these fish indicated coho and chinook were in poorer condition in 1983 than in non-El Niño years. Because adult coho salmon returned to hatcheries at a smaller size, the fecundity (eggs per female) in 1983 was reduced from the 1978–1982 average by 24% at coastal hatcheries and by 27% at Columbia River hatcheries. The fecundity of chinook salmon was unchanged at most hatcheries.     

The vertical distribution of juvenile salmon (Oncorhynchus spp.) and associated fishes in the Columbia River plume

Simultaneous trawling at surface and at depth at one location off the Columbia River, Oregon, in June 2000 identified the depth distribution of juvenile salmonids and associated fishes. Juvenile salmon off the Columbia River were distributed primarily near the surface, within the upper 12 m. Highest densities of subyearling chinook salmon (Oncorhynchus tshawytscha) off the Columbia River were associated with high surface currents and decreasing tidal levels, with time of day possibly a co‐factor. Densities of yearling chinook salmon increased with higher turbidity. Pacific herring (Clupea pallasi) was the most abundant and commonly caught forage fish, with density increasing at night, probably related to diel vertical migration. Catches of juvenile salmonids were not associated with catches of forage fishes. Daytime surface trawling appears to be an appropriate method for assessing the distribution and abundance of juvenile salmonids in marine habitats.     

Sea lice infestation rates on wild and escaped farmed Atlantic salmon (Salmo salar L.) entering the Magaguadavic River, New Brunswick

A monitoring program for the prevalence and intensity of sea lice infestations of wild and escaped farmed salmon has been underway on the Magaguadavic River since 1992. Fish are screened in a fish ladder trap located in freshwater a short distance above the head of tide. No trends with time were evident in observed sea lice burdens, and in all years the majority of salmon, both wild and escapees, had no or low levels of infestation with sea lice. In the spring of 2002, 23 landlocked salmon moving to sea from the Magaguadavic River were acoustically tagged. Two fish returned to the river after a brief period of residence in Passamaquoddy Bay, with significant dermal damage from sea lice. These fish were tracked to areas close to commercial salmon farms.     

Evaluation of an experimental Aeromonas salmonicida epidemic in chinook salmon, Oncorhynchus tshawytscha (Walbaum)

To determine the dynamics of the transmission of Aeromonas salmonicida ssp. salmonicida infection, chinook salmon, Oncorhynchus tshawytscha, were exposed to bacteria by cohabitation. The latent period (time between exposure and infectivity) was determined by exposing a group of chinook salmonid fingerlings to A. salmonicida by bath, then, at daily intervals, by holding five exposed (donor) fish with approximately 50 naive fish for 24 h. The latent period was 3 days post-infection and the time period between the initial exposure to bacteria and the beginning of bacterial shedding was 4.5 days for the same animals. The prevalence and intensity of infection in the donor fish, to which recipient fish were exposed, i.e. the level of exposure, was highly correlated with the development of disease in recipient (susceptible) chinook salmon (r2 = 0.57). An experiment was conducted to determine the daily progress of infection and development of a furunculosis epidemic among recipient fish by cohabiting a single exposed fish with 43 unexposed salmon. At daily intervals, all fish (in seven treatment tanks and one control tank daily) were sacrificed and tested for the presence of A. salmonicida in the kidney (n = 3520). Over 10 days, mean prevalence among recipient fish reached 75% and disease related mortality exceeded 50%. Bacterial concentrations in the water continued to increase over the duration of the experiment in concert with the number of infected animals present in the population.     

Effects of dietary canola oil level on growth, fatty acid composition and osmoregulatory ability of juvenile fall chinook salmon (Oncorhynchus tshawytscha)

This study assessed refined canola oil (CO) as a supplemental dietary lipid source for juvenile fall chinook salmon, Oncorhynchus tshawytscha, parr with respect to possible effects on their growth and osmoregulatory performance and body composition. Diets with equal protein ( 57%) and lipid ( 19%) content (dry weight basis) were supplemented with lipid from either anchovy oil (AO) or CO with AO so that CO comprised 0 (0CO), 11% (11CO), 22% (22CO), 33% (33CO), 43% (43CO) or 54% (54CO) of the dietary lipid content. Triplicate groups of juvenile chinook salmon were fed their prescribed diets for 104 days in freshwater (FW) and 31 days in seawater (SW) after a 4-day transition period. Dietary fatty acid compositions reflected the different proportions of AO and CO in the supplemental lipid. Diet treatment had no effect on fish growth, feed intake, feed efficiency, protein utilization, fish mortality or terminal whole body water and ash percentages. Whole body lipid percentages were higher in 11CO and 43CO fish than in 33CO fish and in 11CO fish versus 22CO fish. Whole body protein percentages were highest in 33CO, 43CO and 54CO fish and lowest in 0CO and 22CO fish. Terminal whole body fatty acid compositions were influenced strongly by the dietary fatty acid compositions. Haematocrit and muscle water percentages were not affected consistently and plasma Na⁺ and Cl⁻ concentrations were unaffected by diet treatment in FW or 24-h seawater challenges during FW residency. Also, diet treatment had no effect on the physiological parameters after SW residency. We conclude that dietary treatment had no effect on fish growth performance under our experimental conditions. Also, the dietary inclusion of CO neither facilitated nor impaired the transfer of chinook salmon parr to seawater. Thus, CO was found to be an excellent and cost-effective source of supplemental dietary lipid for culture of juvenile fall chinook salmon during freshwater residency.     

Parasites and hepatic lesions among pink salmon, Oncorhynchus gorbuscha (Walbaum), during early seawater residence

Juvenile pink salmon, Oncorhynchus gorbuscha (Walbaum), in the Broughton Archipelago region of western Canada were surveyed over 2 years for sea lice (Lepeophtheirus salmonis and Caligus clemensi), gross and microscopic lesions and evidence of infections with viruses and bacteria. The 1071 fish examined had an approximate ocean residence time no longer than 3 months. A high prevalence of degenerative liver lesions, renal myxosporean parasites and a low prevalence of skin lesions and sea lice were observed. No indications of viral or bacterial diseases were detected in either year. The monthly prevalence of sea lice in 2007 (18-51%) was higher than in 2008 (1-26%), and the infestation density exceeded the lethal threshold in only two fish. Degenerative hepatic lesions and renal myxosporean parasites occurred in approximately 40% of the pink salmon examined in June of both years, and the peak monthly prevalence of hepatocellular hydropic degeneration was greater in 2007 (32%, in May) than in 2008 (12%, in June). Logistic regression analysis found skin lesions and hepatocellular hydropic degeneration significantly associated with sea lice. Most parasites and lesions occurred during both years, but the prevalence was often higher in 2007. Fish weight was 35% less in June 2007 than in June 2008, but condition factor was not different. Further research is required to monitor inter-annual variations and aetiology of the liver lesions and to assess their potential role on pink salmon survival.     

Temperature-mediated en route migration mortality and travel rates of endangered Snake River sockeye salmon

Abstract  Conservation efforts for endangered Snake River sockeye salmon ( Oncorhynchus nerka ) have been hindered by high en route adult mortality during their ∼1450 km freshwater spawning migration. Identifying causal factors for this mortality has been difficult given very small (often <10 fish) annual returns in recent decades. However, several hundred hatchery-bred fish returned in 2000 and we intercepted and radio-tagged 31 in mid-migration to monitor behaviours and survival. All fish initially migrated at similar rates, but later-timed fish eventually slowed migration and were far more likely to be unsuccessful. Late-season mortality was strongly associated with water temperatures near tolerance thresholds (21–24 °C). The data also suggest increased risk for fish in poor initial condition (i.e., with injuries or parasites) and probable recent selection against late-timed salmon. Results parallel temperature- and condition-related adult mortality in Columbia and Fraser River sockeye salmon populations and demonstrate the potential vulnerability of marginal southern populations to regional climate warming.     

Infection of gill and kidney of Fraser River sockeye salmon,Oncorhynchus nerka (Walbaum), by Parvicapsula minibicornis and its effect on host physiology

Adult sockeye salmon, Oncorhynchus nerka (Walbaum), migrating upstream in the Fraser River, British Columbia, are exposed to the myxozoan parasite Parvicapsula minibicornis when they enter the river from the ocean. Infections are initially localized in the kidney but have recently been associated with branchitis in one population. Adult fish from five locations in the watershed were sampled to determine whether branchitis was widespread. P. minibicornis infections in kidney glomeruli were prevalent in all samples except for a sample of fish that had just entered the Fraser River from the ocean. For fish captured in spawning streams, parasites were observed in the renal tubules and gill, and branchitis was observed in 70% of fish. Plasma osmolality was negatively correlated with the number of parasites in the kidney tubules, which we hypothesize to be caused by the breach of glomerular membranes as the parasite leaves the fish. Plasma lactate values increased with increasing levels of pathology in gills. These findings support the hypothesis that P. minibicornis impacts the physiology of migrating fish, which may in turn affect the likelihood that adults will be able to migrate and spawn successfully.     

The effect of whole body lipid on early sexual maturation of 1+ age male chinook salmon (Oncorhynchus tshawytscha)

Early sexual maturation of male chinook salmon (maturation 1 to 4 years prior to females in the same age class) results in reduced effectiveness of stock enhancement programs and a financial loss to the salmon farming industry. Previous studies in Atlantic salmon have shown that the age of maturity in males is affected by growth and/or body energy stores, but the relative roles of these two factors are not well understood. Therefore, an experiment was designed to determine when spermatogenesis was initiated, to characterize the endocrine changes during the onset of puberty in male salmon, and to determine if the level of whole-body lipid affects the incidence of early male maturation in a wild stock (Yakima River) of 1+ spring chinook salmon. Fry were fed a commercial diet from February until August and were then divided into groups of 320 fish (mean weight, 5.6 g) and fed one of five experimental diets (two replicate groups/diet) containing 4%, 9%, 14%, 18% or 22% lipid and 82%, 77%, 73%, 69%, or 65% protein for 13 months. Fish were reared on natural photoperiod and ambient temperature (6°C to 16°C), and pair-fed to a level based on the tank with the lowest feed consumption. Fish were weighed monthly and sampled to determine body composition, pituitary follicle-stimulating hormone (FSH) and luteinizing hormone (LH) levels, plasma insulin-like growth factor I (IGF-I) levels, and stage of gonadal development.

Throughout the experimental period the mean fish weight was similar among treatment groups. However, from December through the end of the experiment in the following September, maturing males were significantly larger than nonmaturing fish. Initial lipid levels in 0-age experimental fish were near 6%, which is similar to wild fish of the same stock and age captured in the Yakima River during August. Fish fed diets containing more than 4% lipid increased in whole-body lipid content during the first 2 months of feeding and then maintained at relatively constant levels during the course of the experiment. Whole-body lipid levels for the dietary treatment groups averaged 5.6%, 7.1%, 8.2%, 9.4%, and 9.6% from October through the following September.

Based on histological examination of the testes of experimental fish, type B spermatogonia and primary spermatocytes were first observed in some of the yearling males during November. These were designated maturing males. Pituitary FSH levels were significantly higher in maturing than nonmaturing males at this time and for the remainder of the study. Pituitary FSH levels increased as spermatogenesis proceeded in maturing fish, whereas pituitary LH levels increased in maturing 1+ males only during July and August, when testes were in late stages of spermatogenesis and in September during spermiation. Plasma IGF-I levels were significantly higher in maturing males than nonmaturing fish from December through the end of experiment. Since maturing males were significantly larger than nonmaturing fish of both sexes from December through September, the difference in IGF-I levels could be due to differences in growth or due to maturation.

The percentage of maturing males was significantly influenced by whole-body lipid, increasing from 34% in fish fed the 4% lipid diet to 45% in fish fed the 22% lipid diet. These data suggest that whole-body lipid levels influenced the incidence of maturation of male spring chinook salmon. In addition, both endocrine and histological indicators suggest that maturation was initiated in males approximately a full year prior to the time the fish will spawn.     

Survival of smaller sport caught chinook, Oncorhynchus tshawytscha (Walbaum), and coho, Oncorhynchus kisutch (Walbaum), salmon from Lake Michigan and its management implications

Abstract The survival of small-sized (<50.8 cm) chinook salmon, Oncorhynchus tshawytscha (Walbaum), and coho salmon, Oncorhynchus kitsutch (Walbaum), caught by sport fishing was determined to assess the potential for increasing the size limit for these fish. Fishermen were recently catching smaller salmon than in the 1970s, but salmon growth rates had not changed. To be an effective management option, the survival rate of hooked and returned fish must be high. The overall survival rates were high: 76% for chinook salmon and 70% for coho salmon. There was no significant difference in survival of the coho salmon with size of hook used ( P = 0.31). Any mortality among fish was generally acute; fish hooked deep in the mouth or gills generally bled and died shortly after capture. Fish hooked in the gills had a significantly greater mortality ( P = 0.0002). The overall high survival rate for these species was the result of a small proportion of fish being hooked in the gills or deep in the mouth. Since the survival rate of the salmon was high, the size limit could be increased to allow smaller fish to grow to sizes preferred by sport fishermen.     

Differences in Ichthyophonus prevalence and infection severity between upper Yukon River and Tanana River chinook salmon, Oncorhynchus tshawytscha (Walbaum), stocks

Two genetically distinct populations of chinook salmon, Oncorhynchus tshawytscha (Walbaum), were simultaneously sampled at the confluence of the Yukon and Tanana rivers in 2003. Upper Yukon-Canadian fish had significantly higher infection prevalence as well as more severe infections (higher parasite density in heart tissue) than the lower Yukon-Tanana River fish. Both populations had migrated the same distance from the mouth of the Yukon River at the time of sampling but had significantly different distances remaining to swim before reaching their respective spawning grounds. Multiple working hypotheses are proposed to explain the differences between the two stocks: (1) the two genetically distinct populations have different inherent resistance to infection, (2) genetically influenced differences in feeding behaviour resulted in temporal and/or spatial differences in exposure, (3) physiological differences resulting from different degrees of sexual maturity influenced the course of disease, and (4) the most severely infected Tanana River fish either died en route or fatigued and were unable to complete their migration to the Tanana River, thus leaving a population of apparently healthier fish.     

Chronic glomerulonephritis in returning fall chinook salmon, Oncorhynchus tshawytscha (Walbaum), from the Rogue River, Oregon

Epizootic mortalities of returning adult Rogue River fall chinook salmon, Oncorhynchus tshawytscha (Walbaum), were first observed in 1978. These mortalities reached approximately 76% in 1979 and declined to 30--50% in 1980 and less than 10% in 1981 and 1982. Histopathological examination of kidneys from moribund returns in 1981 and 1982 showed a diffuse chronic glomerulonephritis (GN) in 83% and 30% of the fish examined, respectively. In most cases glomerular damage was severe enough to cause morbidity from osmoregulatory failure. The lesion was markedly similar to immune complex-mediated glomerular disease in higher vertebrates, but tissues for further ultrastructural examination and diagnosis were not processed. Giemsa-stained sections also revealed unidentified sporozoan-like vegetative stages encysted within the extracellular mesangial matrix of glomeruli in both GN and normal kidneys. Although probably incidental to the glomerular lesion, the causal relationship of the organism to GN remains unknown until further investigation. Glomeruli of outmigrating Rogue River fall chinook salmon smolts collected in 1983 were normal and free of the organism.     

Plasma insulin-like growth factor-I concentrations in yearling chinook salmon (Oncorhynchus tshawytscha) migrating from the Snake River Basin, USA

During the parr-to-smolt transformation (smoltification) of juvenile salmonids, preadaptive changes in osmoregulatory and ionoregulatory ability are regulated in part by the growth hormone (GH)/insulin-like growth factor-I (IGF-I) axis. If food intake is sufficient, plasma IGF-I increases during smoltification. On the other hand, plasma IGF-I typically decreases in fasting fish and other vertebrate animals. Because food availability is limited for juvenile salmonids undertaking an extended 6- to 12-week springmigration to and through the Snake-Columbia River hydropower system (northwestern USA), IGF-I concentrations might be expected to decrease, potentially compromising seawater tolerance. To address this possibility,yearling chinook salmon Oncorhynchus tshawytscha reared in three Snake River Basin hatcheries were sampled before release and at two downstream dams. Dry masses ofmigrating fish either did not increase during themigration (in 2000, an average-flow year), or decreased significantly (in 2001, a low-flow year). In both years, plasma IGF-I levels were significantly higher (1.6-fold in 2000, 3.7-fold in 2001) for fish sampled at the last dam on the lower Columbia River than for fish sampled prior to release. Plasma IGF-I concentrations inmigrating fish may, nonetheless, have been nutritionally down-regulated to some degree, because plasma IGF-I concentrations in juvenile chinook salmon captured at a Snake River dam and transported to the laboratory increased in fed groups, but decreased in unfed groups. The ability ofmigrating smolts to maintain relatively elevated IGF-I levels despiteRestricted food intake and loss of body mass is likely related to smoltification-associated changes in hormonal balance.