Changes in GnRH levels in the brain and pituitary gland during migrations of sockeye salmon and chum salmon

Levels of two moleculer types of gonadotropin-releasing hormone (GnRH), salmon GnRH (sGnRH) and chicken GnRH–II (cGnRH–II) in the various brain regions and pituitary gland of sockeye salmon (Oncorhynchus nerka) and chum salmon (O. keta) during smoltification and spawning migration, respectively, were measured using specific time-resolved fluoroimmunoassay (TR-FIA) systems. Changes in sGnRH levels in different brain regions tended to be specifically synchronized with serum thyroid hormone or pituitary gonadotropin (GTH) levels during smoltification and spawning migration, respectively. In contrast, cGnRH–II levels did not show such synchronized changes. SGnRH and cGnRH–II in various brain regions might have different roles during smoltification and spawning migration of salmonid fishes.     

Bilirubin-binding protein in the serum of spawning-migrating chum salmon,Oncorhynchus keta: Its identity with carotenoid-carrying lipoprotein

The serum carotenoid level gradually increased during the spawning migration of chum salmon (Oncorhynchus keta). Zeaxanthin, a yellowish carotenoid, was detected in the serum of spent males, while astaxanthin was a main carotenoid in the serum of males and females at other developmental stages. A high bilirubin level was found in the serum of spent males; the serum of spent male was yellow in colour due to the presence of zeaxanthin and bilirubin, although the serum of the other fish was bright orange.Three types of carotenoid-carrying lipoprotein such as low density lipoprotein, high density lipoprotein (HDL) and very high density lipoprotein fractions were present in the serum. Carotenoid-carrying lipoprotein from the HDL fraction became a main component during spawning migration. The serum bilirubin from the spent male salmon was distributed in the HDL fraction. Because bilirubin, as well as carotenoids, were present in the HDL fraction, the carotenoid-carrying lipoprotein from the HDL fraction was identical with bilirubin-binding protein in the serum of spent males. This carrier lipoprotein had two subunits (molecular weight 24,000 and 12,000); no disulfide bond was detected between the two subunits.     

Capture versus tagging impacts on chum salmon freshwater spawning migration travel times

The spawning migration travel times of chum salmon, Oncorhynchus keta (Walbaum), fitted with gastrically implanted radio tags vs external spaghetti tags were tested for a short [≈60 river km (rkm)] and long migration route (≈730 rkm) on the Koyukuk River, Alaska, USA. Using a novel application of statistical arrival curve models to infer travel times for uncaptured fish, migrations by chum salmon not directly handled during the study were also assessed. Results demonstrated negligible differences in travel times within migration routes between fish fitted only with spaghetti tags and fish fitted with radio tags, indicating low impacts on migration travel behaviour associated with gastric tags once deployed. Conversely, travel times for unhandled fish as inferred by statistical arrival models may have been 12%–24% shorter than those for fish captured with gillnets for tagging. These results suggest that, if present, chum salmon migration behaviour impacts may be more strongly associated with fish capture than tag deployment.     

An evaluation of the potential influence of SST and currents on the oceanic migration of juvenile and immature chum salmon (Oncorhynchus keta) by a simulation model

Using a salmon migration model based on the assumption that swimming orientation is temperature dependent, we investigated the determining factors of the migration of juvenile and immature chum salmon (Oncorhynchus keta) in the North Pacific. We compared the predictions of the model with catch data of immature and juvenile chum salmon collected by Japanese research vessels from 1972 to 1999. The salmon migration model reproduced the observed distributions of immature chum salmon and indicates that passive transport by wind‐driven and geostrophic currents plays an important role in the eastward migration of Asian salmon. These factors result in a non‐symmetric distribution of Asian and North American chum salmon in the open ocean. The directional swimming component contributes to the northward migration in summer. The model results indicate that during the first winter Asian chum salmon swim northward against the southward wind‐driven currents to stay in the western North Pacific. This suggests that Asian chum salmon require more energy to migrate than other stocks during the first winter of their ocean life.     

Immunoreactive somatolactin cells in the pituitary of young,migrating, spawning and spent chinook salmon,Oncorhynchus tshawytscha

Immunocytochemical techniques using an antiserum to cod somatolactin (SL) demonstrated the presence of SL cells in the intermediate lobe of the pituitary in Oncorhynchus tshawytscha. The cells were small in yearling fish. Two groups of maturing fish were studied. In the spring run salmon collected in April and May during the upstream migration, the SL cells appeared stimulated. In September, during spawning, SL cell stimulation was maximal with indices of hypertrophy and degranulation often more marked in females than in males. In the other group, salmon of the fall run collected in the Pacific Ocean in August had well developed gonads, large gonadotropes and abundant SL cells. In spawning salmon (September) the SL cells were stimulated, mainly in females. However, the final stimulation was less intense than in spring run spawning fish. The SL cells were smaller, without evident granule release, but still abundant in spent salmon of the fall run caught at the end of November. Various factors (time spent in rivers before spawning, starvation, decalcification, stress, hypothalamic influences) were considered which might explain differences between spring and fall run salmon. These observations suggest that SL may play a role in the control of gonadal maturation in chinook salmon as it may also do in sockeye and chum salmon previously studied, and that SL cells may be sensitive to the ambient salinity.     

Metabolic adaptations of oxidative muscle during spawning migration in the Atlantic salmon <Emphasis Type="Italic">Salmo salar</Emphasis> L.

The adaptability/plasticity of the highly oxidative red muscle in Atlantic salmon was demonstrated during spawning migration. Substrate concentrations and the enzymatic pathways of ATP production were examined in red muscle obtained from Atlantic salmon at different sites along their migratory route in the Exploits River, Newfoundland, Canada. Individuals were chronologically sampled from a seawater site, two sites upstream, and at spawning. The 20% decrease in salmon body weight during the later stages of migration was accompanied by large decreases (mg dry weight⁻¹) in both glycogen (P < 0.01) and total muscle lipid (P < 0.01). In contrast, water content and protein concentration (mg dry weight⁻¹) of the red muscle increased by 25 and 34%, respectively, at spawning. Enzymes of the glycolytic pathways demonstrated a significant (P < 0.001) decrease in maximal activity as migration proceeded whereas enzymes of the oxidative phosphorylation pathways, specifically the citric acid cycle enzymes, exhibited an increase (P < 0.001) in maximal activity at spawning. The antioxidant enzyme superoxide dismutase also demonstrated an increase (P < 0.001) in maximal activity during the latter stages of migration. These adaptations imply that the red epaxial muscle of Atlantic salmon has a more efficient means of oxidizing lipids, while minimizing free radical damage, during the later stages of migration and spawning, thereby potentially increasing post spawning survival.     

Migratory behaviour of adult wild and escaped farmed Atlantic salmon, Salmo salar L., before, during and after spawning in a Norwegian river

The migratory behaviour of adult wild and escaped farmed Atlantic salmon, Salmo salar L., before, during after spawning in the River Namsen, Norway, was analysed using radio telemetry. The fish were caught, radio tagged and released into the fjord between 7 and 25 km from the river mouth. A significantly higher proportion of wild (74%) than farmed (43%) salmon was subsequently recorded in the river. Wild salmon (33%) were more frequently captured in the sea and in rivers than farmed salmon (14%). The migration speed from release to passing a data logger 11 km upstream from the river mouth was not significantly different between wild (20.6 km day^?1) and farmed (19.8 km day^?1) salmon. Wild salmon tagged when water flow in the river was increasing had a significantly higher migration speed than wild salmon tagged when water flow was decreasing. This was not true for farmed salmon. Farmed salmon were distributed significantly higher up the river than wild salmon during spawning, although both types of fish were found together in spawning areas. Thus, there was no geographical isolation to prevent spawning between wild and escaped farmed salmon. Farmed salmon had significantly more and longer up- and downstream movements than wild salmon during the spawning period. Unlike farmed salmon, the number of riverine movements by wild salmon increased significantly when variation in water flow increased. A smaller proportion of wild (9%) than farmed (77%) salmon survived through the winter after spawning.     

Prespawning migratory behaviour of wild and farmed Atlantic salmon, Salmo salar L, in a north Norwegian river

The migration patterns of wild and released farmed Atlantic salmon, Salmo salar L., were studied by radio-telemetry during migration from entering a river to after spawning. The wild salmon were caught during return migration in bagnets and the farmed salmon were produced in a near by fish farm. Both groups were tagged and released at sea near the river mouth of the River Alta, North Norway. A pronounced individual variation in migration pattern was observed. No significant differences was found between wild and farmed salmon in the distance from entering the river mouth to the place of the first stop (recorded in the same pool for 5 days or longer) and the days from entering the river mouth to arrival at the first stop. For both wild and farmed salmon, mean migration speed was 2.6 km day¹, varying from 0.4 to 11.0 km day^-1. A larger proportion of farmed salmon distributed to the upper part of the river at spawning; mean distances from the river mouth were 30.1 and 19.1 km. respectively. Farmed salmon spent a significantly longer time from entering the river to reaching the area occupied during spawning.     

Effect of food amount and temperature on growth rate and aerobic scope of juvenile chum salmon

Offshore migration of Pacific salmon Oncorhynchus spp. is partly triggered by increasing body size and high motility in the early stages of life. The survival of juvenile salmon may depend on their growth rate during the first few months in the sea, and this factor partly regulates the dynamics of adult populations. Here, we assessed the effects of water temperature and food availability on the growth of juvenile chum salmon O. keta. In addition, by combining the measurements of metabolic performance for growth and activity (Absolute Aerobic Scope: AAS) with a bioenergetics model, we estimated the energy allocation for different activities in the juveniles. Under high temperatures (14 °C), juveniles reared at low food levels (1% body weight) allocated less than half their energy for growth than those reared at high food levels (4% body weight). These findings suggest that high temperature and low food level constrain the growth of juveniles, providing an insight into the effect of the recent increase in warm and low-nutrient water masses on survival of juveniles and catches of adult chum salmon on the Pacific side of Honshu Island, Japan.

     

Stability in reproductive timing and habitat usage of Chinook salmon across six years of varying environmental conditions and abundance

The spatial and temporal distribution of spawning activity by autumn‐run Chinook salmon Oncorhynchus tshawytscha (Walbaum) was examined across multiple years. The study period included two years of extreme drought conditions when water temperatures in the spawning reach of the Stanislaus River were considered sub‐optimal for spawning and egg incubation. Despite varying levels of superimposition, redd counts and densities remained stable and positively associated with river location, indicating that superimposition may be driven by habitat preference rather than the absence of suitable spawning locations. Spawning occurred slightly later during drought years (6–10 days later compared with wetter years). This delay was attributable to deferred migration instead of deferred spawning, and the majority of redds were constructed at water temperatures exceeding the optimal temperature range. As a consequence, estimates of juvenile production during 2014 and 2015 were among the lowest on record. These findings may be related in part to the high hatchery contribution to the population, above‐average temperatures during spawning and incubation, and superimposition rates. Management recommendations include adequate cold‐water storage in the upstream reservoir, refined spawning habitat restoration techniques in the light of superimposition rates and, on a broader system scale, actions that reduce the amount of stray hatchery Chinook salmon.     

Changes in plasma somatolactin levels during spawning migration of chum salmon (Oncorhynchus keta)

Plasma somatolactin (SL) concentrations were examined in chum salmon in relation to gonadal maturation; immature salmon in the Bering Sea at various stages of maturation, and mature salmon during upstream migration caught at the ocean, bay and river. Plasma SL concentrations as well as plasma prolactin (PRL) and growth hormone (GH) levels in the immature fish caught in the Bering Sea were maintained essentially at similar levels. Plasma SL in mature salmon increased significantly from the fish in the ocean to the fish in the river in both sexes. Although all the fish had fully developed gonads, females completed ovulation while still in the bay, whereas final spermeation in males was achieved after entry into the river. Thus, no clear correlation was seen between plasma SL levels and final gonadal maturation. On the other hand, plasma PRL concentrations in both male and female fish were higher in the fish in the river than those in the ocean and bay, and plasma GH levels were higher in both sexes in the fish in the bay and river than those in the ocean. Plasma levels of triglycerides, glucose, free fatty acids and ionized sodium and calcium were also examined. Significant-negative correlations were seen between plasma SL and plasma ionized calcium in mature male salmon, and between plasma SL and plasma triglycerides in mature female salmon. Although our findings do not rule out the possibility of the involvement of SL in final maturation, the results indicate that SL seems to be involved at least in energy and/or calcium metabolism during the spawning migration.     

Monitoring the spawning behaviour of chum salmon with an acceleration data logger

Abstract –  The continuous observation of salmon behaviour in some wild environments can be extremely difficult. We recorded spawning behaviour of female chum salmon ( Oncorhynchus keta ) in the field simultaneously using visual observation and fish-borne data loggers with two-axis accelerometer sensors. Using only acceleration records, behaviours were successfully classified into the eight well-known components of spawning behaviour: swimming, nosing, exploratory digging, nest digging, probing, oviposition, covering and post-spawning digging. To understand how the female chum salmon modulates spawning behaviours in relation to changes in environmental conditions, we compared the behaviours of salmon during normal flow of clear water to those of salmon during the heavy flow of turbid water after a storm. Salmon in the normal flow showed all eight behaviours, whereas salmon in the heavy flow showed only three behaviours: swimming, nosing and exploratory digging. The proportion of time spent on swimming was greater in the heavy flow than in the normal flow (mean of 98.47% vs. 92.84%). Moreover, the proportion of tail beating in swimming was greater in the heavy flow (77.86%) than in the normal flow (15.63%). Our results indicate that the behaviour of female chum salmon was strongly affected by the heavy flow of turbid water after a storm. The recording of accelerations is a promising method for clarifying the spawning behaviour of salmonids in the wild where continuous visual observation is too difficult.     

Effect of global warming on the life history and population dynamics of Japanese chum salmon

We have reviewed the effects of long-term climatic/oceanic conditions on the growth, survival, production dynamics, and distribution of Hokkaido chum salmon Oncorhynchus keta in Japan during the period 1945–2005 using path analysis, back-calculation, and scale analyses, and applied a prediction method based on the SRES-A1B scenario of the intergovernmental panel on climate change. The populations of Hokkaido chum salmon were found to have had high growth rates at age 1 year since the late 1980s. Path analysis indicated that the growth at age 1 year in the Okhotsk Sea was directly affected by warm sea surface temperature associated with global warming, with the increased growth at age 1 year resulting in higher rates of survival and large population sizes. Predictions on the global warming effects on the chum salmon were (1) decreased carrying capacity and distribution area, (2) occurrence of a strong density-dependent effect, and (3) loss of migration route to the Sea of Okhotsk, especially for Hokkaido chum salmon. We have also outlined the future challenges of establishing a sustainable conservation management scheme for salmon that include adaptive management and precautionary principles, as well as conservation of natural spawning populations and recovery of natural river ecosystems in Japan despite the warming climate.     

Mechanism of body cavity temperature regulation of chum salmon (Oncorhynchus keta) during homing migration in the North Pacific Ocean

Vertical movement patterns of five chum salmon (Oncorhynchus keta) during homing migration were examined using archival tags. The standard deviation of the depth and ambient and body cavity temperatures during daytime were larger than those during night‐time. Vertical movements through the thermocline with a periodicity of less than 1 h were observed during daytime in addition to the diel vertical movement patterns in the open ocean. During these periods of frequent short‐term vertical movements, the difference between the body cavity temperature and ambient temperature was large while the variance of the body cavity temperature was less than that of the ambient temperature. From the results of a random simulation, the variation of the body cavity temperature was shown to decrease due to these periodic high frequency movements in comparison with random vertical movements. The whole‐body heat‐transfer coefficient k (s^?1), which was estimated by a heat budget model, was 1.48 × 10^?3. The k of chum salmon was larger than that of bigeye tuna (Thunnus obesus) by about one order of magnitude for the cooling of the body. The k of chum salmon did not change like tuna, which are physiologically adapted to conserve body cavity temperature. This indicates that the regulation of body cavity temperature by chum salmon is dependent on the vertical movements only. The maintenance of the body cavity temperature is concluded to be advantageous for their maturation and growth from the relationship between energy input and output during their homing migration.     

Changes in rhodopsin-porphyropsin ratio of chum and pink salmon

ABSTRACT:   In order to investigate the changes in rhodopsin-porphyropsin ratio of chum salmon and pink salmon in relation to the change in their habitat, the ratios were analyzed in individual fish prior to stocking, during the sea run, homing, and upstream migration. The ratio in both the species gradually increased during the sea run. However, the ratio decreased after upstream river migration. Moreover, in the sea, the rhodopsin-porphyropsin ratio of chum salmon was always slightly higher than that of pink salmon. The largest difference in the individual variation was observed in the individuals caught with a set net placed near the mouth of a home river.     

Current status of natural spawning of chum salmon <Emphasis Type="Italic">Oncorhynchus keta</Emphasis> in rivers with or without hatchery stocking on the Japan Sea side of northern Honshu,Japan

Information on the status of natural spawning is needed on the Japan Sea side of northern Honshu, Japan for ecosystem-based sustainable management of chum salmon resources. We conducted on-site visual surveys in October–December of 2015 and 2016 that targeted spawning chum salmon redds in all rivers?>?5 km long (total 94 rivers) in Akita, Yamagata, Niigata (including Sado Island), and Toyama prefectures. The ratio of rivers found to host natural reproduction to the total number of surveyed rivers was 93.6% (44/47) in stocked rivers and 74.5% (35/47) in non-stocked rivers. These results show that there is a wide occurrence of natural reproduction of chum salmon in these rivers, regardless of the history of hatchery stocking. The density of spawning redds (number of redds/1000 m²) as an indicator of chum salmon escapements did not differ (P?=?0.54) between stocked rivers (mean 3.5, N ?=?49) and non-stocked rivers (mean 2.4, N? =?36),when rivers where no redds were observed were excluded from the analysis. These results suggest that chum salmon escapements into non-stocked rivers may not be negligible. Conservation measures for wild fish are needed in stocked and non-stocked rivers to promote enhancement programs based on natural reproduction.     

Density‐dependent effects of eastern Kamchatka pink salmon (Oncorhynchus gorbuscha) and Japanese chum salmon (O. keta) on age‐specific growth of western Alaska chum salmon

Hatcheries release >4.5 billion juvenile Pacific salmon (Oncorhynchus spp.) into the North Pacific Ocean annually, raising concerns about competition with wild salmon populations. We used retrospective scale analysis to investigate how the growth of chum salmon (O. keta) from western Alaska is affected by the abundance of chum salmon from Japanese hatcheries and wild pink salmon (O. gorbuscha) from the Russian Far East. Over nearly five decades, the growth of Kuskokwim River chum salmon was negatively correlated with the abundance of Japanese hatchery chum salmon after accounting for the effects of sex and spring/summer sea‐surface temperature in the Bering Sea. An effect of wild eastern Kamchatka pink salmon abundance on the growth of Kuskokwim River salmon was detectable but modest compared to the intraspecific competitive effect. A decrease in Japanese hatchery chum salmon releases in 2011–2013 was not associated with increased growth of Bering Sea chum salmon. However, the abundance of wild chum salmon from the Russian Far East increased during that time, possibly obscuring reduced competition with hatchery chum salmon. Our results support previous evidence that chum salmon are affected by intraspecific competition, and to a lesser extent interspecific competition, in the North Pacific, underscoring that the effects of salmon hatchery production transcend national boundaries.     

Grey mullet roe contains yolk protein having IgE cross-reactivity to chum salmon roe major allergen (Onc k 5)

The major allergenic protein in grey mullet roe was immunologically characterized using rabbit IgG against the β′-component, which is a vitellogenin fragment and the major allergen in chum salmon roe, and the IgE cross-reactivity between salmon and mullet roe was examined using chum salmon roe-allergic patients' sera. Grey mullet β′-component was identified in the yolk protein and consisted of 16 and 30 kDa proteins that showed marked reaction with the anti-β′-component IgG and shared N-terminal amino acid sequence similarity with vitellogenin. The grey mullet β′-component consisted of six different isoforms generated from vitellogenin A and vitellogenin B, and they inhibited the IgE-binding reaction between chum salmon β′-component and a chum salmon roe-allergic patient’s serum. Thus, grey mullet roe may be an allergenic seafood capable of introducing IgE cross-reactivity to chum salmon roe through the β′-component.     

Seasonal variations in and effect of incubation water temperature on vertebral number in naturally spawning chum salmon <Emphasis Type="Italic">Oncorhynchus keta</Emphasis>

Seasonal variations and reaction norms for vertebral number (V _N) in response to incubation water temperature were estimated in adult and juvenile naturally spawning chum salmon Oncorhynchus keta. The mean V _N of adults varied according to spawning time; the early-spawning population had higher V _N values than the late-spawning population. Moreover, the mean V _N values in the early-spawning population decreased with seasonal changes, whereas V _N values in the late-spawning population remained stable. Chum salmon embryos in three full-sib families were incubated at five different temperatures until hatching, and the V _N values of the resulting juveniles were analyzed. The V _N reaction norm to incubation water temperature showed a V-shaped curve that was lowest at an intermediate temperature. The mean V _N at the same incubation temperature varied among the three families. These results suggest that V _N values in chum salmon are influenced by genetic components and incubation water temperatures. V _N may be a useful parameter for estimating the environmental conditions during ontogenesis and the genetic background by detecting population changes.     

Population structure and stock identification of chum salmon Oncorhynchus keta from Japan determined by microsatellite DNA variation

Variation at 14 microsatellite loci was surveyed in 26 chum salmon Oncorhynchus keta populations from Japan, one population from West Kamchatka and three populations from North America to determine population structure. Microsatellites were then applied to estimate stock composition of chum salmon in mixed-stock fisheries. The genetic differentiation index (F _st) over all populations and loci was 0.031, with individual locus values ranging from 0.010 to 0.081. Seven regional populations were observed in Japanese chum salmon, with late-run populations from the Pacific Coast of Honshu the most distinct. Japanese populations displayed greater genetic diversity than did those in North America. Transplantation history in some Japanese river populations influenced their present genetic characteristics. Analysis of simulated mixtures from fishery sampling suggested that accurate and precise regional estimates of stock composition should be produced when the microsatellites were used to estimate stock compositions. Stock compositions for a 2005 sample of maturing, migrating chum salmon off the north-west coast of Hokkaido near the border of the Sea of Japan and the Sea of Okhotsk indicated that this region may be a migration corridor for Hokkaido populations from the Sea of Japan coast. Microsatellites have the ability to provide fine-scale resolution of stock composition in Japanese coastal fisheries.