Changes in habitat availability for outmigrating juvenile salmon (Oncorhynchus spp.) following estuary restoration

The restoration of the Nisqually River Delta (Washington, U.S.A.) represents one of the largest efforts toward reestablishing the ecosystem function and resilience of modified habitat in the Puget Sound, particularly for anadromous salmonid species. The opportunity for outmigrating salmon to access and benefit from the expansion of available tidal habitat can be quantified by several physical attributes, which are related to the ecological and physiological responses of juvenile salmon. We monitored a variety of physical parameters to measure changes in opportunity potential from historic, pre‐restoration, and post‐restoration habitat conditions at several sites across the delta. These parameters included channel morphology, water quality, tidal elevation, and landscape connectivity. We conducted fish catch surveys across the delta to determine if salmon was utilizing restored estuary habitat. Overall major channel area increased 42% and major channel length increased 131% from pre‐ to post‐restoration conditions. Furthermore, the results of our tidal inundation model indicated that major channels were accessible up to 75% of the time, as opposed to 30% pre‐restoration. Outmigrating salmon utilized this newly accessible habitat as quickly as 1 year post‐restoration. The presence of salmon in restored tidal channels confirmed rapid post‐restoration increases in opportunity potential on the delta despite habitat quality differences between restored and reference sites.     

Enhanced invertebrate prey production following estuarine restoration supports foraging for multiple species of juvenile salmonids (Oncorhynchus spp.)

Estuaries provide crucial foraging resources and nursery habitat for threatened populations of anadromous salmon. As such, there has been a global undertaking to restore habitat and tidal processes in modified estuaries. The foraging capacity of these ecosystems to support various species of out‐migrating juvenile salmon can be quantified by monitoring benthic, terrestrial, and pelagic invertebrate prey communities. Here, we present notable trends in the availability of invertebrate prey at several sites within a restoring large river delta in Puget Sound, Washington, U.S.A. Three years after the system was returned to tidal influence, we observed substantial additions to amphipod, copepod, and cumacean abundances in newly accessible marsh channels (from 0 to roughly 5,000–75,000 individuals/m²). In the restoration area, terrestrial invertebrate colonization was dependent upon vegetative cover, with dipteran and hymenopteran biomass increasing 3‐fold between 1 and 3 years post‐restoration. While the overall biodiversity within the restoration area was lower than in the reference marsh, estimated biomass was comparable to or greater than that found within the other study sites. This additional prey biomass likely provided foraging benefits for juvenile Chinook, chum, and coho salmon. Primary physical drivers differed for benthic, terrestrial, and pelagic invertebrates, and these invertebrate communities are expected to respond differentially depending on organic matter exchange and vegetative colonization. Restoring estuaries may take decades to meet certain success criteria, but our study demonstrates rapid enhancements in foraging resources understood to be used for estuary‐dependent wildlife.     

Gauging resource exploitation by juvenile Chinook salmon (Oncorhynchus tshawytscha) in restoring estuarine habitat

In the context of delta restoration and its impact on salmonid rearing, success is best evaluated based on whether out‐migrating juvenile salmon can access and benefit from suitable estuarine habitat. Here, we integrated 3 years of post‐restoration monitoring data including habitat availability, invertebrate prey biomass, and juvenile Chinook salmon (Oncorhynchus tshawytscha) physiological condition to determine whether individuals profited from the addition of 364 ha of delta habitat in South Puget Sound, Washington, United States. Productivity in the restored mudflat was comparable to reference sites 3 years after dike removal, surpassing a mean total of 6 million kJ energy from invertebrate prey. This resulted from the development of a complex network of tidal channels and a resurgence in dipteran biomass that was unique to the restoration area. Consequently, a notable shift in invertebrate consumption occurred between 2010 and 2011, whereby individuals switched from eating primarily amphipods to dipteran flies; however, dietary similarity to the surrounding habitat did not change from year to year, suggesting that this shift was a result of a change in the surrounding prey communities. Growth rates did not differ between restored and reference sites, but catch weight was positively correlated with prey biomass, where greater prey productivity appeared to offset potential density‐dependent effects. These results demonstrate how the realized function of restoring estuarine habitat is functionally dependent. High prey productivity in areas with greater connectivity may support healthy juvenile salmon that are more likely to reach the critical size class for offshore survival.     

Timing games in the reproductive phenology of female pacific salmon (Oncorhynchus spp.)

We use a game-theoretic framework to investigate the reproductive phenology of female kokanee (Oncorhynchus nerka). As in the other semelparous species of Pacific salmon, females construct nests in gravel, spawn with males, bury their fertilized eggs, and defend their nest sites until they die several days later. Later-breeding females may reuse previous nest sites, and their digging behavior is thought to subject previously buried eggs to mortality. Using game-theoretic models, we show that females can reduce this risk by allocating resources to longevity (the period between arrival and death) as opposed to eggs. Waiting before territory settlement is also expected if it allows females to conserve energy and delay senescence. The models demonstrate how these costs and benefits interact to select for a seasonal decline in longevity, a well-known phenomenon in the salmonid literature, and a seasonal decline in wait duration. Both of these predictions were supported in a field study of kokanee. Female state of reproductive maturity was the most important proximate factor causing variation in longevity and wait duration. With more than 30% of territories being reused, dig-up is likely an important selective force in this population.     

Chemosensory responses of juvenile Coho salmon, Oncorhynchus kisutch, Dolly Varden, Salvelinus malma, and sculpins (Cottus spp.) to eggs and other tissues from adult Pacific salmon

The carcasses of semelparous Pacific salmon (Oncorhynchus spp.) provide nutrients that enter aquatic ecosystems by various pathways, including direct consumption of tissue by fishes. Salmonids and other species frequently eat eggs and other tissues from dead salmon but the roles of vision and olfaction are unclear, as is the relative attraction to different tissues. Accordingly, we conducted a series of in situ experiments using minnow traps in two natural streams in Alaska to test the relative roles of chemosensory and visual cues in attraction of fishes to eggs from adult Pacific salmon, and then compared catch rates of traps baited with eggs, muscle, liver, and testis. Experiments indicated that chemical traces were necessary and sufficient to attract juvenile Dolly Varden (Salvelinus malma), coho salmon (Oncorhynchus kisutch), and sculpins (Cottus spp.) into traps. Combining both sites, 70 salmonids and 19 sculpins were trapped using visual and chemical cues, and 53 and 21, respectively, for traps with only chemical cues. Traps with only the sight of eggs caught no salmonids and only 5 sculpins, comparable to empty control traps. In addition, eggs were markedly more attractive than the other tissues, trapping 68?% of the salmonids and 69?% of the sculpins, compared to 14?% and 15?% for muscle tissue, 12?% and 11?% for liver, and 6?% and 5?% for the testis. Visual cues undoubtedly play a role in egg consumption in streams, but these experiments indicated a very important role of chemical traces in attracting fish to the vicinity of the eggs, and selective attraction of eggs over other salmon tissues.     

Detection of infection and susceptibility of different Pacific salmon stocks (Oncorhynchus spp.) to the haemoflagellate Cryptobia salmositica

The haematocrit centrifugation technique, modified by keeping the haematocrit tubes cold (between 1 and 10 C), was sensitive for detecting light infections of Cryptobia salmositica (as few as 75 flagellates per ml of blood). In wet mount preparations, infections lighter than 7.5 X 10(3) flagellates per ml of blood could not be detected consistently. Different Pacific salmon stocks from British Columbia demonstrated differences in susceptibility to C. salmositica in experimental studies using laboratory reared juvenile fish. Oncorhynchus keta and Oncorhynchus tshawytscha from the Big Qualicum River stocks (Vancouver Island), and Oncorhynchus nerka from the Fulton River stock (Skeena River system), were all equally susceptible and suffered high mortalities at low exposures (100 flagellates in 0.1 ml physiological saline inoculated intraperitoneally per fish). Oncorhynchus nerka from the Weaver Creek stock (Fraser River system) was the most resistant with no mortalities even at exposures of 10(6) flagellates (in 0.1 ml physiological saline) per fish. Oncorhynchus kisutch seemed to be slightly less resistant than the Weaver Creek O. nerka, but fewer than 16% of the inoculated fish died. Oncorhynchus kisutch from the Big Qualicum River seemed to be slightly more resistant than O. kisutch from the Capilano River stock (a coastal river near Vancouver), with fewer mortalities and lighter infections when the experiments were terminated. Differences in susceptibility are believed to be associated with innate, genetically transmitted resistance.     

Morpho-ecological patterns and feeding of coho salmon Oncorhynchus kisutch fry in the period of downstream migration

The principal patterns of feeding of downstream-migrant and coastal young of the year of coho salmon are considered. The so-called potential downstream migrants and residents, which differ in size and feeding patterns, were distinguished among coastal fry. It is assumed that identification of particular coastal specimens to phenotypic groups of downstream migrants or nonmigrants of young of the year of coho salmon may be determined even in the daytime by such parameters as the body length, feeding intensity, and food spectrum.     

Spatial and temporal patterns of growth and consumption by juvenile spring/summer Chinook salmon Oncorhynchus tshawytscha

Somatic growth is often used as a metric of habitat quality, but such an approach has limitations because growth results from complex interactions between abiotic and biotic factors. In this study, we derived estimates of weekly growth (based on otolith growth increments) across several months for four populations of threatened Chinook salmon, Oncorhynchus tshawytscha, from the Salmon River Basin, Idaho, USA. Although mean stream temperature varied by 2–5 °C across populations, growth across the season did not vary significantly by population. To investigate this further, we applied a bioenergetics model that produced estimates of consumption rates. We then examined how growth and consumption rates varied according to rearing stream and day in the season. Using generalized least squares models, somatic growth (g?day^?1) was best explained by stream and date, yet a model with only date had moderate support, and thus indicated limited support for stream effects. Specific consumption rate (g?g^?1?day^?1 and J?g^?1?day^?1) was best explained using a model that included main effects of stream and date. These findings are consistent with the hypothesis that higher temperatures confer higher metabolic costs that require greater consumption to produce similar growth rates in cooler streams. This highlights that similarity in growth rate among streams may mask changes in individual behavior and/or energetic acquisition associated with differences in temperatures among streams. Results of this study represent the first steps towards identifying factors that underlie important population level and habitat quality differences.     

Effect of hunger on the daily pattern of feeding rates in juvenile pink salmon, Oncorhynchus gorbuscha Walbaum

Juvenile pink salmon, Oncorhynchus gorbuscha , in two test groups of differing hunger level fed continuously on live copepods, maintained at high densities, throughout 12 h feeding periods in the laboratory. Analysis of video films showed that mean feeding rates were initially (first 10 min) between 51.5 and 63.8 prey-capture attempts (snaps) fish^-1 10 min^-1, but declined abruptly with increasing satiation to relatively constant levels of about 5.0 and 8.0 snaps fish^-1 10 min^-1, which were maintained during the subsequent 11 h of feeding. The data suggests that after the initial filling of their stomachs with food, juvenile pink salmon keep their stomachs full by feeding at a rate that balances the gastric evacuation rate of 18.6 mg prey h^-1 at 11° C. Less than 15% of the fish's stomach contents need be evacuated apparently for spontaneous feeding to resume or occur. Increased hunger level (by increasing a pre-test food deprivation period from 24 to 72 h) resulted in fish increasing their average feeding rate and thus their ration consumed from 23.6 to 39.8% dry body weight per 12 h.     

Effects of melatonin feeding on smoltification in masu salmon (Oncorhynchus masou)

Influences of photoperiod on plasma melatonin profiles and effects of melatonin administration on long-day-induced smoltification in masu salmon (Oncorhynchus masou) were investigated in order to reveal the roles of melatonin in the regulation of smoltification in salmonids. Under light-dark (LD) cycles, plasma melatonin levels exhibited daily variation, with higher values during the dark phase than during the light phase. The duration of nocturnal elevation under short photoperiod (LD 8:16) was longer than that under long photoperiod (LD 16:8). Melatonin feeding (0.01, 0.1 and 1 mg/kg body weight) elevated plasma levels of melatonin in a dose-dependent manner for at least 7 h but not for 24 h. When masu salmon reared under short photoperiod were exposed to long photoperiod (LD 16:8) and fed melatonin (1 mg/kg body weight) 7 hours before the onset of darkness, a significantly smaller proportion of smolts appeared in the melatonin-fed group after 32 days than in the control group. However, after 59 days of the treatment, there was no difference in the proportion of smolts between the control and melatonin-treated groups. Thus, melatonin feeding mimicked the effects of short photoperiod, which delays but does not completely suppress smoltification. These results indicate that the day length is transduced into changes in the duration of nocturnal elevation in plasma melatonin levels, and that artificial modification of the plasma melatonin pattern possibly delays the physiological processes of smoltification induced by long-day photoperiodic treatment.     

Diel feeding periodicity, daily ration and prey selection of a riverine population of juvenile chinook salmon, Oncorhynchus tshawytscha (Walbaum)

The diel feeding periodicity, daily ration and prey selection of juvenile chinook salmon, Oncorhynchus tshawytscha , were studied in relation to the available prey. Maximum dry weight of food intake occurred about dawn, when mayflies were the major prey, but the greatest number of freshly eaten prey occurred during the afternoon, when chironomids and terrestrial dipterans predominated. Feeding activity at night was low, with smaller mayflies comprising up to 50% of the prey. During the day the young salmon fed selectively on chironomids and the larger mayflies, while trichopterans and terrestrial taxa were under-represented in the diet. Food consumption over the 24-h period averaged 8.3% of the fish dry body weight. Prey abundance in the drift explained about 50% of the composition of the diet. Although the fish selected larger mayflies, size apparently was not a main criterion for selection because chironomids, although smaller than mayflies, were also frequently eaten. Previous dietary experience of the fish and the diel pattern of prey abundance appear to best explain the selective feeding of juvenile chinook salmon.     

Disruption of seasonality in growth hormone-transgenic coho salmon (Oncorhynchus kisutch) and the role of cholecystokinin in seasonal feeding behavior

Seasonal variation in daily food intake is a well-documented phenomenon in many organisms including wild-type coho salmon where the appetite is noticeably reduced during periods of decreased day length and low water temperature. This reduction may in part be explained by altered production of cholecystokinin (CCK) and growth hormone (GH). CCK is a hormone produced in the brain and gut that mediates a feeling of satiety and thus has an inhibitory effect on food intake and foraging behaviour. Growth hormone (GH) enhances feeding behaviour and consequently growth, but its production is reduced during winter. The objectives of this study were: first, to compare the seasonal feeding behaviour of wild and GH-transgenic coho salmon; second, to determine the behavioural effect of blocking the action of CCK (by using devazepide) on the seasonal food intake; and third, to measure CCK expression in brain and gut tissues between the two genotypes across seasons. We found that, in contrast to wild salmon, food intake in transgenic salmon was not reduced during winter indicating that seasonal control of appetite regulation has been disrupted by constitutive production of GH in transgenic animals. Blocking of CCK increased food intake in both genotypes in all seasons. The increase was stronger in wild genotypes than transgenic fish; however blocking CCK in wild-type fish in winter did not elevate appetites to levels observed in the summer. The response to devazepide was generally faster in transgenic than in wild salmon with more rapid effects observed during summer than during winter, possibly due to a higher temperature in summer. Overall, a seasonal effect on CCK mRNA levels was observed in telencephalon with levels during winter being higher compared to the summer in wild fish, but with no seasonal effect in transgenic fish. No differences in seasonal CCK expression were found in hypothalamus. Higher levels of CCK were detected in the gut of both genotypes in winter compared to summer. Thus, CCK appears to mediate food intake among seasons in both wild-type and GH-transgenic salmon, and an altered CCK regulation may be responsible at least in part for the seasonal regulation of food intake.     

Physiological impact of acute molybdenum exposure in juvenile kokanee salmon (Oncorhynchus nerka)

A series of experiments were conducted to determine the physiological impact of acute sublethal molybdenum exposure to juvenile kokanee salmon (Oncorhynchus nerka Kennerlyi). Molybdenum was found to be relatively non-toxic to kokanee as the 96 h LC(50) was greater than 2,000 mg Mo l(-1). Exposure to either 25 or 250 mg Mo x l(-1) for 7 days was found to stimulate a significant 1.6- to 1.7-fold increase in ventilation which was later characterized to be dose-dependent between 5 and 250 mg Mo l(-1). Acute sublethal molybdenum exposure was found to have little or no impact on kokanee oxygen consumption at rest or immediately following a bout of forced activity or on physiological indicators of stress such as plasma lactate, sodium and cortisol. Despite these findings, prior exposure to 25 or 250 mg Mo l(-1) resulted in post-exercise loss of equilibrium and exercise-induced delayed mortality that were not observed in controls. Molybdenum accumulation in gill and liver of kokanee was also characterized. The findings of this study suggest that despite the non-toxic nature of molybdenum, acute sublethal exposure to this metal has physiological consequences to those fish exposed even for only a brief period. Further studies are needed to more fully elucidate the metabolism and mode of action of this metal in fish.     

Diet,feeding patterns,and prey selection of subyearling Atlantic salmon (Salmo salar) and subyearling chinook salmon (Oncorhynchus tshawytscha) in a tributary of Lake Ontario

Since juvenile Atlantic salmon (Salmo salar) and Chinook salmon (Oncorhynchus tshawytscha) occupy a similar habitat in Lake Ontario tributaries, we sought to determine the degree of diet similarity between these species in order to assess the potential for interspecific competition. Atlantic salmon, an historically important but currently extirpated component of the Lake Ontario fish community, are the focus of a bi‐national restoration effort. Presently this effort includes the release of hatchery produced juvenile Atlantic salmon in Lake Ontario tributaries. These same tributaries support substantial numbers of naturally reproduced juvenile Pacific salmonids including Chinook salmon. Subyearling Atlantic salmon and subyearling Chinook salmon had significantly different diets during each of the three time periods examined. Atlantic salmon fed slightly more from the benthos than from the drift and consumed mainly chirononmids (47.0%) and ephemeropterans (21.1%). The diet of subyearling Chinook salmon was more closely associated with the drift and consisted mainly of chironomids (60.2%) and terrestrial invertebrates (16.0%). Low diet similarity between subyearling Atlantic salmon and subyearling Chinook salmon likely minimizes competitive interactions for food between these species in Lake Ontario tributaries. However, the availability of small prey such as chironomids which comprise over 50% of the diet of each species, soon after emergence, could constitute a short term resource limitation. To our knowledge this is the first study of interspecific diet associations between these two important salmonid species.     

Physiological mechanisms of imprinting and homing migration in Pacific salmon Oncorhynchus spp

After several years of feeding at sea, salmonids have an amazing ability to migrate long distances from the open ocean to their natal stream to spawn. Three different research approaches from behavioural to molecular biological studies have been used to elucidate the physiological mechanisms underpinning salmonid imprinting and homing migration. The study was based on four anadromous Pacific salmon Oncorhynchus spp., pink salmon Oncorhynchus gorbuscha, chum salmon Oncorhynchus keta, sockeye salmon Oncorhynchus nerka and masu salmon Oncorhynchus masou, migrating from the North Pacific Ocean to the coast of Hokkaido, Japan, as well as lacustrine O. nerka and O. masou in Lake Toya, Hokkaido, where the lake serves as the model oceanic system. Behavioural studies using biotelemetry techniques showed swimming profiles from the Bering Sea to the coast of Hokkaido in O. keta as well as homing behaviours of lacustrine O. nerka and O. masou in Lake Toya. Endocrinological studies on hormone profiles in the brain-pituitary-gonad axis of O. keta, and lacustrine O. nerka identified the hormonal changes during homing migration. Neurophysiological studies revealed crucial roles of olfactory functions on imprinting and homing during downstream and upstream migration, respectively. These findings are discussed in relation to the physiological mechanisms of imprinting and homing migration in anadromous and lacustrine salmonids.     

Effect of Cryptobia salmositica-induced anorexia on feeding behavior and immune response in juvenile rainbow trout Oncorhynchus mykiss

At 10 degrees C, rainbow trout Oncorhynchus mykiss (n = 13 per group) infected with Cryptobia salmositica Katz, 1951 became anorexic at 3 wk post-infection (w.p.i.), with feed-intake decreasing significantly from 1.33 to 0.94% body weight (b.w.). Anorexia was most severe at 4 w.p.i. (0.80% b.w.), coinciding with peak parasitemia (9.2 x 10(6) parasites ml blood(-1)) and anemia. At 8 w.p.i., fish had recovered their appetite although they still had contained detectable parasites (6.8 x 10(5) parasites ml(-1)) and were anemic (pack cell volume, PCV, of 24.4%). However at 5 degrees C, anorexia occurred at 5 w.p.i. (0.81% b.w.), and was most severe at 7 w.p.i. (0.40% b.w.). At 8 w.p.i. (0.43% b.w.), fish displayed high parasitemia (4.6 x 10(6) parasites ml(-1)) and low PCV (10.8%). Fish at 5 degrees C had lower gastric evacuation (GE) rates (GE48h) than 10 degrees C fish, however there were no differences between infected and naive fish at both temperatures. Before anorexia, there was no significant correlation between mean share of meal (MSM, a measure of how food was partitioned within a group) and coefficient of variation in feeding but this became significant during anorexia (p = 0.02 and p = 0.0002 at 10 and 5 degrees C respectively). Significant correlations were detected between b.w. and MSM before onset of anorexia at 10 degrees C (p = 0.005) and 5 degrees C (p = 0.02); this was maintained at 10 degrees C (p = 0.001) but not at 5 degrees C (p = 0.98). Fish on an anorexic diet (0.93% b.w.) responded well at 10 degrees C to a live C. salmositica vaccine; this could partly be due to constant antigenic stimulation by the live vaccine.     

Understanding the behavior and ecology of Chinook salmon (Oncorhynchus tshawytscha) on an important feeding ground in the Gulf of Alaska

Population declines and demographic changes of Chinook salmon (Oncorhynchus tshawytscha), have been documented throughout this species’ range, though information on natural and anthropogenic mechanisms related to these changes are not fully understood. To provide insights into marine behaviors and survival of Chinook salmon, 40 pop-up satellite archival tags (PSATs), that collected environmental data, were attached to large (69–100 cm FL) Chinook salmon caught in the marine waters of Cook Inlet, Alaska. PSATs provided evidence of predation on tagged Chinook salmon by ectothermic and unconfirmed predators, and provided valuable information about the migratory characteristics and occupied depths and temperatures of this species while occupying Cook Inlet and the Gulf of Alaska. The results from this study suggest that late-marine mortality of Chinook salmon of a variety of stock-origins by apex predators is more common in Cook Inlet than previously thought, and may be used to improve our understanding this species’ population dynamics. Furthermore, results from this study adds to the existing knowledge of marine habitat use by Chinook Salmon and may be useful in assessing the vulnerability and interactions between this species and anthropogenic activities.