Iteroparity in complex mating systems of steelhead Oncorhynchus mykiss (Walbaum)

This study investigated diverse reproductive types in complex mating systems of steelhead Oncorhynchus mykiss . Postspawned steelhead (kelts) were sampled during attempted downstream migration over Lower Granite Dam on the Snake River, U.S.A. Multilocus microsatellite genotypes (14 loci) were used to assign unknown origin, kelt individuals to upstream populations of origin. Results indicated that iteroparity is a life-history trait that remains in several tributaries of the Snake River basin despite strong selection against downstream adult passage because of hydroelectric dams. The largest populations of steelhead in the Snake River, however, were only weakly represented (Clearwater River = 7·5% and Salmon River = 9·4%, respectively) in the kelt steelhead mixture relative to the Grande Ronde River (18·2%), Imnaha River (17·4%), Pahsimeroi Hatchery (25·2%) and Asotin Creek (22·2%). A lack of correlation between population escapement size and kelt proportions ( P > 0·05) suggests that iteroparity was not uniformly expressed across populations, but was significantly negatively correlated with body size ( P < 0·05). Iteroparity may be a valuable source of genetic variability and a conservation priority, especially in years with poor recruitment or in recently bottlenecked populations.     

Post-spawn migrations of hatchery-origin Oncorhynchus mykiss kelts in the Central Valley of California

We used acoustic telemetry to study the post-spawn movement of Oncorhynchus mykiss kelts released in April 2005 and 2006 from the Coleman National Fish Hatchery, Anderson, CA. Following release, O. mykiss kelts demonstrated both anadromous and non-anadromous life histories, with some fish alternating life history strategies between years. Anadromy was most common, characterized by a short-term residence near the release site, followed by sustained downstream emigration once initiated. O. mykiss kelts demonstrating anadromy arrived at the Golden Gate Bridge from April to mid-July. Repeat spawning migrations of anadromous O. mykiss kelts began from late-September through October of the year of release. High fidelity back to Battle Creek was observed, occurring from late-September through November. While most O. mykiss kelts were anadromous, at least 10 % remained in freshwater, or residualized. O. mykiss kelts that residualized demonstrated two distinct patterns of movement: 1) residency near the release location, and 2) potamodromy. Overall survival was high with 36 % and 48 % of O. mykiss kelts released making a repeat spawning migration and demonstrating iteroparity in 2005 and 2006, respectively. Increase in body lengths of O. mykiss kelts that returned to the Coleman NFH were significantly greater for anadromous fish, compared to fish that residualized, but survival was higher for fish that residualized. Release of hatchery O. mykiss kelts could result in both positive and negative genetic and ecological effects to hatchery- and naturally-producing salmonids. We believe the benefits of releasing O. mykiss kelts at the Coleman NFH, including increased numbers and size of fish in the recreational fishery and genetic and demographic benefits to the hatchery brood stock outweigh the limited risk to natural populations that would result from predation and competition of the relatively small number of O. mykiss kelts that resided in fresh water.     

Marine feeding of anadromous Salmo trutta during winter

Juvenile and adult anadromous trout Salmo trutta utilize the sea for feeding during the winter in the Skagerrak. This finding conflicts with the traditional view that anadromous trout overwinter in fresh water. Adults, just prior to spawning, were captured at sea in October to December, and spent fish were caught at sea from October to April, showing that the fish may leave the stream and move to sea just after spawning and spend the winter there. During mid‐winter (January to February), the feeding probability (chance of finding a fish with food in its stomach) increased markedly with increasing body length, with no similar effect during early and late winter (October to December and March to April). Among individuals with food in their stomach (72·5%), there was no evidence for variation in feeding intensity [stomach fullness = (mass of stomach content)(fish body mass)⁻¹] among early, mid‐, and late winter.     

Life-history effects of migratory costs in anadromous brown trout

Mean size of sexually mature anadromous brown trout (sea trout) Salmo trutta in south-east Norway increased significantly with migratory distance ( D ) between the feeding area at sea and the spawning area in fresh water, from 32 cm for those spawning close to the river mouth to 43 cm at the spawning grounds 40 km inland. This was largely due to an increased size of the smallest anadromous spawners with increasing D . The raised mean size of the long-distance migrants is paralleled by an increased age at sexual maturity. Body mass at the same length of sea trout decreased with D in fresh water, meaning that the fish moving far inland was slimmer than those spawning near the coast. Gonadal mass of first-time spawning anadromous males declined significantly with D , and the fecundity and the ratio of fecundity over mean mass of the individual eggs adjusted for variation in fish mass, increased with D . There was no clear relationship between the ratio of anadromous to resident fish and D , probably because more variables than D , influence this relationship in the study streams.     

Who are the missing parents? Grandparentage analysis identifies multiple sources of gene flow into a wild population

In order to increase the size of declining salmonid populations, supplementation programmes intentionally release fish raised in hatcheries into the wild. Because hatchery-born fish often have lower fitness than wild-born fish, estimating rates of gene flow from hatcheries into wild populations is essential for predicting the fitness cost to wild populations. Steelhead trout (Oncorhynchus mykiss) have both freshwater resident and anadromous (ocean-going) life history forms, known as rainbow trout and steelhead, respectively. Juvenile hatchery steelhead that 'residualize' (become residents rather than go to sea as intended) provide a previously unmeasured route for gene flow from hatchery into wild populations. We apply a combination of parentage and grandparentage methods to a three-generation pedigree of steelhead from the Hood River, Oregon, to identify the missing parents of anadromous fish. For fish with only one anadromous parent, 83% were identified as having a resident father while 17% were identified as having a resident mother. Additionally, we documented that resident hatchery males produced more offspring with wild anadromous females than with hatchery anadromous females. One explanation is the high fitness cost associated with matings between two hatchery fish. After accounting for all of the possible matings involving steelhead, we find that only 1% of steelhead genes come from residualized hatchery fish, while 20% of steelhead genes come from wild residents. A further 23% of anadromous steelhead genes come from matings between two resident parents. If these matings mirror the proportion of matings between residualized hatchery fish and anadromous partners, then closer to 40% of all steelhead genes come from wild trout each generation. These results suggest that wild resident fish contribute substantially to endangered steelhead 'populations' and highlight the need for conservation and management efforts to fully account for interconnected Oncorhynchus mykiss life histories.     

Comparative sensitivities of diagnostic procedures used to detect bacterial kidney disease in salmonid fishes

Kidney and spleen homogenates from each of 60 coho salmon (Oncorhynchus kisutch) and steelhead trout (Salmo gairdneri) were examined for detection of Renibacterium salmoninarum. The proportions of positives differed widely with the detection procedures used: in coho salmon, 5% were positive by the Gram-stain procedure, 10% by the direct fluorescent antibody test, 48% by bacteriological isolation, 65% by staphylococcal coagglutination, and 73% by counterimmunoelectrophoresis; in steelhead trout, 3% were positive by Gram-stain, 8.3% by fluorescent antibody, 17% by bacteriological isolation, and 67% by counterimmunoelectrophoresis. Renibacterium salmoninarum was not detected in either coho salmon or steelhead trout by immunodiffusion analysis.     

Anadromy and the marine migrations of Pacific salmon and trout: Rounsefell revisited

Anadromy is a defining trait in salmonid fishes but it is expressed to different extents among the species in the family, as reviewed in a classic paper by Rounsefell (1958). The present paper re-examines the subject, assessing the degree of anadromy within the genus Oncorhynchus, using Rounsefell’s six criteria: extent of migrations at sea, duration of stay at sea, state of maturity attained at sea, spawning habits and habitats, post-spawning mortality, and occurrence of freshwater forms of the species. The genus ranges from pink salmon (O. gorbuscha), the most fully anadromous species in the family, to entirely non-anadromous species closely related to rainbow trout (O. mykiss), including Mexican golden trout (O. chrysogaster), Gila and Apache trout (O. gilae), and sub-species of cutthroat trout (O. clarki). This paper provides updated information on anadromy and marine migration patterns, emphasizing the iteroparous species, cutthroat (O. clarki) and rainbow (O. mykiss) trout. These two species display widely ranging patterns of anadromy, including truly “landlocked” populations and residents with easy access to the sea. Anadromous rainbow trout (known as steelhead) populations also vary greatly in their distribution at sea, incidence of repeat spawning, and associated traits. We conclude, as did Rounsefell, that anadromy is not a single trait with two conditions (anadromous or non-anadromous). Rather, it reflects a suite of life history traits that are expressed as points along continua for each species and population. Further research is needed in the marine ecology of all species but especially trout, as they are less well known but apparently more variable in patterns of anadromy and life history than salmon species.     

Adaptive trade-offs in juvenile salmonid metabolism associated with habitat partitioning between coho salmon and steelhead trout in coastal streams

1. Adaptive trade-offs are fundamental to the evolution of diversity and the coexistence of similar taxa and occur when complimentary combinations of traits maximize efficiency of resource exploitation or survival at different points on environmental gradients. 2. Standard metabolic rate (SMR) is a key physiological trait that reflects adaptations to baseline metabolic performance, whereas active metabolism reflects adaptations to variable metabolic output associated with performance related to foraging, predator avoidance, aggressive interactions or migratory movements. Benefits of high SMR and active metabolism may change along a resource (productivity) gradient, indicating that a trade-off exists among active metabolism, resting metabolism and energy intake. 3. We measured and compared SMR, maximal metabolic rate (MMR), aerobic scope (AS), swim performance (UCrit) and growth of juvenile hatchery and wild steelhead and coho salmon held on high- and low-food rations in order to better understand the potential significance of variation in SMR to growth, differentiation between species, and patterns of habitat use along a productivity gradient. 4. We found that differences in SMR, MMR, AS, swim performance and growth rate between steelhead trout and coho salmon were reduced in hatchery-reared fish compared with wild fish. Wild steelhead had a higher MMR, AS, swim performance and growth rate than wild coho, but adaptations between species do not appear to involve differences in SMR or to trade-off increased growth rate against lower swim performance, as commonly observed for high-growth strains. Instead, we hypothesize that wild steelhead may be trading off higher growth rate for lower food consumption efficiency, similar to strategies adopted by anadromous vs. resident brook trout and Atlantic salmon vs. brook trout. This highlights potential differences in food consumption and digestion strategies as cryptic adaptations ecologically differentiating salmonid species. 5. We hypothesize that divergent digestive strategies, which are common and well documented among terrestrial vertebrates, may be an important but overlooked aspect of adaptive strategies of juvenile salmonids, and fish in general.     

Maturation and spawning processes of anadromous and resident pond smelt in lake ogawara

Pond smelt,Hypomesus nipponensis McAllister, in Lake Ogawara demonstrate alternative life history strategies, as evidenced by the coexistence of anadromous and resident fish. However, it is unknown if anadromous and resident groups interbreed. In this study, maturation and spawning processes were examined and compared between anadromous and resident groups. Histological observations indicated negligible variation in the maturational stage composition of oocytes, the frequency of oocyte diameter being unimodal for all specimens at different maturational stages. Oocytes were absent in the ovaries of spent fish. Accordingly, the species can be considered a semelparous spawner with unimodal oocyte diameter distribution. Temporal changes in the proportion of spent fish were compared between anadromous and resident groups. Spawning of both groups began in late March and peaked over April 8–12. Although both groups did not differ significantly in the period of peak spawning, anadromous fish finished spawning earlier than resident ones. Anadromous fish were not able to spawn upon migration into Lake Ogawara, and quickly matured after immigration, contrasting with resident fish.     

Temperature effects on juvenile anadromous salmonids in California’s central valley: what don’t we know?

The anadromous Chinook salmon (Oncorhynchus tshawytscha) (4 runs) and steelhead (rainbow trout, O. mykiss), are both native to Californias Sacramento-San Joaquin River (SSJR) system, whose watershed encompasses the central valley of California. The SSJR system holds the southernmost extant Chinook salmon populations in the Eastern Pacific Ocean, whereas coastal anadromous steelhead populations are found at more southerly latitudes. Populations of both species of anadromous salmonid have experienced dramatic declines during the past 100 years, at least partly from water impoundments and diversions on most central valley rivers and their tributaries. These changes restricted the longitudinal distribution of these salmonids, often forcing the superimposition of steelhead populations and Chinook salmon populations in the same reaches. This superimposition is problematic in part because the alterations to the river systems have not only changed the historic flow regimes, but have also changed the thermal regimes, resulting in thermally-coupled changes in fish development, growth, health, distribution, and survival. Given the highly regulated nature of the system, resource managers are constantly trying to strike a balance between maintaining or increasing the population size of anadromous fish runs and with other demands for the water, such as irrigation and water quality. To do so, in this review, we summarize the published information on the temperature tolerance and growth of the stream-associated life stages of these two valuable species, which are so central to the natural heritage of the State and its cultures. We show that many of these limits and growth-related effects are specific regarding life stage and that some may be specific to distinct strains or races of Chinook salmon and steelhead within the system. Because the number of published studies on the physiology of central valley salmonids was surprisingly low, we also use this review to highlight critical areas where further research is needed. Overall, this review should assist biologists and resource decision-makers with improved understanding for the protection and enhancement of these native fishes.     

Influence of aperiodic summer droughts on leaf litter breakdown and macroinvertebrate assemblages: testing the <Emphasis Type="Italic">drying memory</Emphasis> in a Central Apennines River (Aterno River,Italy)

Management of multiple exploited stocks of anadromous salmonids in large catchments requires understanding of movement and catchment use by the migrating fish and of their harvesting. The spawning migration of sea trout (Salmo trutta) and Atlantic salmon (Salmo salar) was studied in the River Tweed, UK, using acoustic telemetry to complement exploitation rate data and to quantify catchment penetration. Salmon (n = 79) and sea trout (n = 65) were tagged in the tidal-influenced Tweed in summer–autumn. No tagged salmon left the river before spawning, but 3% (2010) and 8% (2011) of pre-spawning sea trout dropped out. Combined tag regurgitation/fish mortality in salmon was 12.5%, while trout mortality was 6% (2010) and 0% (2011). The estimated spawning positions of salmon and sea trout differed; tagged salmon were mostly in the main channel while trout occurred mostly in the upper Tweed and tributaries. Early fish migrated upstream slower than later fish, but sea trout moved through the lower-middle river more quickly than salmon, partly supporting the hypothesis that the lower exploitation rate in autumn of trout (1 vs 3.3% for salmon) there is generated by differences in migration behaviour.     

Age, sex ratio and timing of the catch of kelts and ascending Atlantic salmon in the subarctic River Teno

By 15 June, 82% of the catch of Atlantic salmon Salmo salar kelts had been taken from the middle part of River Teno, northern Scandinavia. The median date of capture was 4 June for males and 8 June for females. Salmon of 1–4 sea–winters (SW) of both sexes survived spawning to return to sea as kelts. Among males, 1 SW kelts were caught earliest in the spring and 3 SW latest, but among females 4 SW were earliest, then 3 SW and finally 1 and 2 SW. There were 17 river and sea–age combinations among the kelts compared with 23 among the ascending salmon. The smolt age distribution and the mean smolt age differed significantly only between female 2 SW ascending salmon (3·97 years) and kelts (4·14 years). The proportion of 1 SW females was higher and that of 3 SW males lower among kelts than among ascending salmon. The proportion of males among 1 SW ascending salmon was 80% but among kelts only 57%. Similarly, the proportion of males among 3 SW fish was 21% for ascending salmon but only 7% for kelts. Hence overwinter mortality was higher among males. Male and female kelts of 1 and female kelts of 2 SWhad a greater mean length than ascending salmon in corresponding groups indicating a better survival of larger fish within an age group. Grilse ascend rivers after most kelts have left, but the main catch of ascending 2–3 SW salmon takes place concurrently with kelts leaving the river, inadvertently targeting kelts in the fishery.     

The post‐spawning movements of migratory brown trout Salmo trutta L.

The post spawning behaviour of sea trout Salmo trutta was studied over a 2 year period in the river and estuary of the River Fowey, south‐west England. Forty‐five sea trout kelts were trapped immediately after spawning in December and intraperitoneally tagged with miniature acoustic transmitters. The subsequent emigration into coastal waters was monitored using acoustic receivers deployed throughout the river catchment. The levels of gill Na⁺K⁺ATPase activity in sea trout kelts sampled at the same time as the tagged fish were within the range of 2·5 to 4·5 μmol Pi per mg protein per h indicating that the post‐spawning fish were not physiologically adapted to salt water. The tagged kelts were resident in fresh water between 4 and 70 days before entering the estuary. Sixty two per cent of the tagged kelts subsequently migrated successfully into coastal waters, with a higher success rate for male fish (75%) than females (58%). There was a significant size related difference in the run‐timing of the kelts with the larger fish moving more quickly into coastal waters after spawning than smaller fish. Seaward migration within fresh water was predominantly nocturnal and generally occurred in conjunction with increasing river discharge and rising water temperature. Migration through the estuary continued to be predominantly nocturnal and occurred during an ebbing tide. Residency within the estuary varied amongst individuals although it was invariably short, with most fish moving out into coastal waters within one to two tidal cycles. Five tagged kelts returned from the coastal zone and re‐entered fresh water during April and June. Marine residence time varied between 89 and 145 days (mean 118 days) and the minimum estimated marine survival was c. 18%. One of these sea trout was subsequently recaptured after successfully spawning in the vicinity where it had been previously tagged demonstrating a degree of spawning site fidelity.     

Large‐scale parentage analysis reveals reproductive patterns and heritability of spawn timing in a hatchery population of steelhead (Oncorhynchus mykiss)

Understanding life history traits is an important first step in formulating effective conservation and management strategies. The use of artificial propagation and supplementation as such a strategy can have numerous effects on the supplemented natural populations and minimizing life history divergence is crucial in minimizing these effects. Here, we use single nucleotide polymorphism (SNP) genotypes for large‐scale parentage analysis and pedigree reconstruction in a hatchery population of steelhead, the anadromous form of rainbow trout. Nearly complete sampling of the broodstock for several consecutive years in two hatchery programmes allowed inference about multiple aspects of life history. Reconstruction of cohort age distribution revealed a strong component of fish that spawn at 2 years of age, in contrast to programme goals and distinct from naturally spawning steelhead in the region, which raises a significant conservation concern. The first estimates of variance in family size for steelhead in this region can be used to calculate effective population size and probabilities of inbreeding, and estimation of iteroparity rate indicates that it is reduced by hatchery production. Finally, correlations between family members in the day of spawning revealed for the first time a strongly heritable component to this important life history trait in steelhead and demonstrated the potential for selection to alter life history traits rapidly in response to changes in environmental conditions. Taken together, these results demonstrate the extraordinary promise of SNP‐based pedigree reconstruction for providing biological inference in high‐fecundity organisms that is not easily achievable with traditional physical tags.     

Primary and secondary oocyte growth dynamics in anadromous semelparous Allis shad Alosa alosa

We analysed the ovarian dynamics of the anadromous semelparous allis shad Alosa alosa for which our working hypothesis was that mature pre-spawning females would have very low or even exhausted primary growth (PG) oocyte reserves; semelparity has been linked with the depletion of the pool of PG oocytes. To test this hypothesis, the PG oocytes were enumerated, their recruitment pattern to the secondary growth (SG) phase was analysed and their potential replenishment from the pool of oogonia was examined in females caught very close to the Mondego River mouth, in central Portugal and along the river. The development of the SG oocytes was also analysed, the fecundity (batch, total and annual) values were estimated and the intensity of atresia was quantified. Ovarian samples and histological sections were investigated in parallel. A dynamic recruitment pattern of PG oocytes to the SG phase was revealed, where all PG oocytes were recruited and were not replenished by oogonia. Annual fecundity was subject to down-regulation due to atresia prior to spawning and its size was multiple times higher than the size of batch fecundity. Lack of population synchronicity in ovarian development and spawning migration was also observed. This multifaceted analysis of the ovarian dynamics of this species will contribute to management efforts for this critically endangered and economically important fish throughout its geographical distribution. The results reported in this study will also assist in unravelling the complexity of the early processes of oogenesis in fish.     

Archival and acoustic tags reveal the post-spawning migrations, diving behavior, and thermal habitat of hatchery-origin Sacramento River steelhead kelts (Oncorhynchus mykiss)

Acoustic and geolocating archival tags were deployed on steelhead kelts to study their post-spawning migrations, diving behavior and thermal habitat. Fourteen reconditioned steelhead kelts were surgically implanted with LAT2510 archival and V7-2 L acoustic tags, and released from Coleman National Fish Hatchery. An array of acoustic receivers in the Sacramento River and Delta, and the San Francisco Bay estuary detected the downstream movement of the steelhead. Two steelhead kelts (J and M) with archival tags were recovered at the hatchery after 219 and 285 days at liberty respectively. Based on changes in geolocations, vertical movements, and water temperatures, the migrations of steelhead J and M were divided into five and six phases respectively. Steelhead J moved into coastal California waters while steelhead M remained in freshwater for the majority of its time at liberty. Large increases in temperature and opacity were recorded before and after the ocean phases, likely as the steelhead moved through the Sacramento Delta. Both steelhead kelts remained relatively close to the surface throughout their migration but there were diurnal differences in the vertical movements. In freshwater, the steelhead tended to be deeper during the day (e.g., steelhead J: 3.08?±?1.50 vs 1.65?±?1.15 m, day vs night) but in the ocean, the steelhead were typically deeper during the night (1.32?±?1.61 vs 5.63?±?6.11 m). Both steelhead kelts appeared to be less oceanic than a previous study in Scott Creek, a small coastal stream approximately 100 km south of the mouth of San Francisco Bay. Although this study has a small sample size, the results suggest that steelhead kelts from a large river and estuary system, like the Sacramento River, may be relatively less oceanic than steelhead kelts from small coastal streams.     

Isolation and characterization of Edwardsiella tarda from fall chinook salmon (Oncorhynchus tshawytscha).

A new bacterial pathogen of chinook salmon (oncorhynchus tshawytscha) was isolated from fish in Oregon's Rogue River. The bacteria are biochemically and serologically related to strains of Edwardsiella tarda. Initially isolated from chinook salmon, the bacteria were also pathogenic for steelhead and rainbow trout (Salmo gairdneri), and channel catfish (Ictalurus punctatus). The 50% lethal doses for chinook salmon, steelhead trout, and channel catfish injected intraperitoneally and maintained in 18 degrees C water were 4.1 x 10(6), 5.6 x 10(6), and 4.0 x 10(5) respectively. When chinook salmon and rainbow trout were injected intraperitoneally and held in 12 degrees C water, the mean lethal doses were 6.4 x 10(7) and 1.7 x 10(6), respectively. The invasiveness of the organism was low in steelhead trout exposed to the bacteria by the waterborne route. The optimum growth temperature of the bacteria in brain heart infusion broth was approximately 35 degrees C. The guanine plus cytosine content of DNA obtained from E. tarda isolated from salmon was 59 mol%.     

Swimming performance of various freshwater Newfoundland salmonids relative to habitat selection and fishway design

Swimming ability of wild brook trout Salvelinus fontinalis , brown trout Salmo trutta , anadromous Atlantic salmon Salmo salar , and landlocked Atlantic salmon was examined using fixed and increasing velocity tests. Although brook trout and salmon parr were collected from the same site, brook trout were found generally in slow-moving pools whereas salmon were more common in faster riffle areas. Salmon parr could hold station indefinitely in currents in which brook trout could only maintain themselves briefly. Therefore, selection of fast-water areas by salmon parr may impose a velocity barrier to sympatric juvenile brook trout, reducing competition between the species. Performance comparisons also indicate that anadromous Atlantic salmon possess slightly greater sustained ability than landlocked salmon, possibly due to altered selective pressure associated with their different life histories. Finally, fishways and culverts in Newfoundland can now be designed using models generated from performance data collected from native salmonid species.     

Variations of life history strategy of the arctic lamprey <Emphasis Type="Italic">Lethenteron camtschaticum</Emphasis> from the Utkholok River (Western Kamchatka)

Morphobiological traits of the lamprey from the Utkholok River (Western Kamchatka) are investigated for the first time. All present forms of lamprey (typically anadromous, anadromous early maturing forma praecox, resident) do not differ by diagnostic characters and belong to one species Lethenteron camtschaticum. Males prevail among mature specimens of various forms, smolts, and ammocoetes at later developmental stages. The forma praecox is represented by males by 92%. A hypothesis is suggested explaining the relationships of the forms and the mechanism of realization of different types of life history strategy in relation to energy resources of water bodies and the kind of feeding of different groups of ammocoetes (organic detritus or Pacific salmon carcasses decomposed after spawning).     

A comparison of cystatin activity in the various tissues of chum salmon Oncorhynchus keta between feeding and spawning migrations

• 1.1. Cystatin was found to be widely distributed in various tissues of chum salmon. Most of the cystatins in the salmon tissues appeared to have a molecular weight between 10,000 and 20,000. They were considered to belong to the low molecular weight form cystatin, the type 1 and/or type 2 cystatins.
• 2.2. The activity in the liver of the salmon in spawning migration was significantly higher than that of the fish in feeding migration, while the activities of the serum, kidney, intestine, stomach, gill, skin and white muscle in spawning migration were apparently lower than that of the fish in feeding migration.
• 3.3. Such differences in the cystatin activity were considered to relate closely to the physiological conditions such as sexual maturation and/or starvation during spawning migration of the fish.