Innate and enhanced predator recognition in hatchery-reared chinook salmon

We used a laboratory behaviour assay to investigate how innate predator recognition, handling stress, retention time, and number of conditioning events might affect chemically mediated anti-predator conditioning for hatchery-reared chinook salmon, Oncorhynchus tshawytscha. Juvenile chinook salmon with no prior exposure to predatory stimuli exhibited innate fright responses to northern pikeminnow, Ptychocheilis oregonensis, odour, regardless of whether the salmon came from a population that exists in sympatry or allopatry with northern pikeminnows. Juvenile chinook salmon exhibited enhanced predator recognition following a single conditioning event with conspecific extract and northern pikeminnow odour. Handling similar to what hatchery salmon might experience prior to release did not substantially reduce the conditioned response. When we conditioned juvenile chinook salmon in hatchery rearing vessels, fish from tanks treated once exhibited a conditioned response to northern pikeminnow odour in aquaria, but only for one behaviour (feeding response), and fish treated twice did not respond. The results suggest that enhanced recognition of predator stimuli occurs quickly, but may be to some extent context-specific, which may limit conditioned fright responses after release into the natural environment.     

Effects of rapid experimental temperature increases on acute physiological stress and behaviour of stream dwelling juvenile chinook salmon

We experimentally heated small streams in summer and investigated the short-term behavioural changes and physiological stress responses of juvenile chinook salmon (Oncorhynchus tshawytscha). We rapidly raised temperatures ∼1–4 °C for 1.5 h above ambient levels of ∼7–15 °C in groundwater fed tributary streams and ∼19–23 °C in side-channel streams. Juvenile chinook rearing in groundwater fed tributaries were generally unaffected behaviourally; however, we found that temperature increase caused fish in the tributary trials to be physiologically stressed (elevations in mean cortisol concentrations ranged from 116% to 253%). Side-channel trials caused some mortality of juvenile chinook and a stronger display of behaviours indicative of stress and avoidance such as erratic swimming, abnormal posture, and aggregative behaviour. Foraging rates increased over 56 times in response to heating in side-channel trials. Cortisol levels did not increase in side-channel trials, but rather showed a trend to levels below control values suggesting an impaired stress response possibly due to chronic stress. Our results may reflect conservative responses in terms of what we may find with other salmonid species since juvenile chinook have been described as the most tolerant of the Pacific salmon species to elevated temperatures.     

Effect of differential gastric evacuation and multispecies prey items on estimates of daily energy intake in juvenile chinook salmon

Synopsis The caloric density of stomach contents in juvenile chinook salmon,Oncorhynchus tshawytscha, was not affected by gastric evacuation, suggesting a constant caloric density of stomach contents during evacuation. Differences in the caloric density of prey consumed did affect caloric density of stomach contents over a 24-h period. Consumption of the amphipodCorophium sp. was associated with reduced caloric densities of stomach contents. During periods whenCorophium contributed more than 4% of the stomach contents, average caloric density declined from 5.56 to 5.33 kcal g^–1. Despite this difference, estimates of daily energy intake of juvenile chinook salmon were only 3%, greater when developed from the mean caloric density of stomach contents excludingCorophium.     

Ontogenetic variations in the type and size of prey consumed by juvenile coho,Oncorhynchus kisutch,and chinook,O. tshawytscha,salmon

Synopsis Stomach contents of juvenile coho,Oncorhynchus kisutch, and chinook,O. tshawytscha, salmon collected in purse seines off the coast of Washington and Oregon were examined for variations related to predator size. There was a general trend toward increasing consumption of fish with increasing body size, due mainly to the increase in northern anchovy biomass consumed by the larger salmon. Most of the major prey taxa showed significant differences among the size classes examined for both salmon species. There was a direct relationship between predator and prey size for both coho and chinook, but considerable variation was found in prey length consumed within each size class. Prey width did not provide as good a fit as prey length for either species. In general, coho consumed larger fish prey in relation to their body length than chinook but there were substantial differences by month or year of collection.     

The effects of turbidity and vegetation on the risk of juvenile salmonids, Oncorhynchus spp., to predation by adult cutthroat trout, O. clarkii

Synopsis During their seaward migration, juvenile salmonids encounter structural and visual cover which varies between and within watersheds. In this study, the effects of two types of cover (turbidity and artificial vegetation) on the predation mortality of juvenile salmonids exposed to fish piscivores was investigated in outdoor concrete ponds. During experiments, adult coastal cutthroat trout, Oncorhynchus clarkii clarkii, were allowed to feed on juvenile salmonid prey — chinook salmon, O. tshawytscha, chum salmon, O. keta, sockeye salmon, O. nerka, and cutthroat trout — in separate trials. Daily instantaneous per capita predation rate was determined for each turbidity and vegetation treatment, within each trial. Mean predation rates varied between 1% and 76% daily. In the presence of cover, mean daily predation rates were 10–75% lower than those in controls (no vegetation and clear water), depending on prey species. Predation rates were significantly lower in the presence of vegetation cover and did not covary with prey size or species. The effects of turbidity were generally not significant and were not additive with the effects of vegetation. However, turbidity appeared to significantly reduce the effectiveness of vegetation as cover for juvenile chinook and sockeye salmon. We suggest that these two forms of cover do not affect risk of predation by fish piscivores to juvenile salmonids via the same mechanism.     

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.     

Exposure of juvenile chinook and chum salmon to chemical contaminants in the Hylebos Waterway of Commencement Bay, Tacoma, Washington

The Hylebos Waterway is an industrialized waterway ofCommencement Bay, Tacoma, Washington, that is severelycontaminated with aromatic and chlorinatedhydrocarbons in the sediment. Juvenile chinook (Oncorhynchus keta) and chum salmon (O.tshawytscha) inhabit this waterway for a few days orweeks during their outmigration from freshwaterstreams to saltwater. The purpose of thisinvestigation was to determine to what degree juvenilechum and chinook salmon captured from the HylebosWaterway might bioaccumulate organic contaminants. These levels of exposure will be compared to previousstudies where such exposures have been linked tobiological dysfunction in juvenile salmon. Theresults showed that juvenile chum and chinook salmonfrom the Hylebos Waterway take up a wide range ofchemical contaminants, compared to fish fromhatcheries or reference estuaries. These contaminantsinclude high and low molecular weight polycyclicaromatic hydrocarbons (PAHs), polychlorinatedbiphenyls (PCBs, including the toxic congeners 105 and118), hexachlorobutadiene (HCBD), hexachlorobenzene(HCB), DDTs, heptachlor, and several pesticides. Immunohistochemical examination of the gill and gut injuvenile chum salmon from the Hylebos Waterway showedthe induction of the P450 metabolizing enzyme. Moreover, concentrations of contaminants in juvenilechinook and chum salmon from the Hylebos Waterway arecomparable to levels previously shown to be associatedwith biological injury in juvenile chinook salmon,such as impaired growth, suppression of immunefunction as demonstrated by reduced B cell function,and increased mortality following pathogen exposure.     

Predators reverse the direction of density dependence for juvenile salmon mortality

The effect of predators on prey populations depends on how predator-caused mortality changes with prey population density. Predators can enforce density-dependent prey mortality and contribute to population stability, but only if they have a positive numerical or behavioral response to increased prey density. Otherwise, predator saturation can result in inversely density-dependent mortality, destabilizing prey populations and increasing extinction risk. Juvenile salmon and trout provide some of the clearest empirical examples of density-dependent mortality in animal populations. However, although juvenile salmon are very vulnerable to predators, the demographic effects of predators on juvenile salmon are unknown. We tested the interactive effects of predators and population density on the mortality of juvenile Atlantic salmon (Salmo salar) using controlled releases of salmon in natural streams. We introduced newly hatched juvenile salmon at three population density treatments in six study streams, half of which contained slimy sculpin (Cottus cognatus), a common generalist predator (18 release sites in total, repeated over two summers). Sculpin reversed the direction of density dependence for juvenile salmon mortality. Salmon mortality was density dependent in streams with no sculpin, but inversely density dependent in streams where sculpin were abundant. Such predator-mediated inverse density dependence is especially problematic for prey populations suppressed by other factors, thereby presenting a fundamental challenge to persistence of rare populations and restoration of extirpated populations.     

Physiological stress responses to confinement in diploid and triploid Atlantic salmon

Confinement of juvenile Atlantic salmon Salmo salar from four different populations (all–female diploids, all–female triploids, mixed sex diploids and mixed sex triploids) either before (FW parr) or after (SW smolts) transfer to sea elevated plasma cortisol and plasma lactate from control levels irrespective of ploidy status. Both before and after confinement, plasma cortisol levels in SW smolts (6–174 ng ml^–1) were higher than those in FW parr (4–58 ng ml^–1), which possibly reflected the physiological challenge of acclimation to SW. Mixed sex populations of SW smolts had higher cortisol levels than all–female populations. The duration of confinement (1, 3 or 6 h) affected the magnitude of the plasma cortisol and lactate responses in SW smolts. Plasma cortisol levels in diploid and triploid SW smolts subjected to 2 h of confinement decreased to pre–stress levels within 6 h post–confinement. Plasma lactate levels were not significantly different from pre–stress levels 48 h after confinement. As no difference exists in primary and secondary stress responses of Atlantic salmon of differing ploidy status, it is unlikely that differences in mortality rates between diploid and triploid populations under commercial conditions can be attributed to differences in their physiological responses to periods of stress lasting up to 6 h.     

Wild chinook salmon survive better than hatchery salmon in a period of poor production

The population dynamics of chinook salmon (Oncorhynchus tshawytscha) from the Cowichan River on Vancouver Island, British Columbia, Canada are used by the Pacific Salmon Commission as an index of the general state of chinook salmon coast wide. In recent years the production declined to very low levels despite the use of a hatchery that was intended to increase production by improving the number of smolts entering the ocean. In 2008, we carried out an extensive study of the early marine survival of the hatchery and wild juvenile chinook salmon. We found that both rearing types mostly remained within the Gulf Islands study area during the period when most of the marine mortality occurred for the hatchery fish. By mid September, approximately 1.3% of all hatchery fish survived, compared to 7.8%–31.5% for wild fish. This six to 24 times difference in survival could negate an estimated increased egg-to-smolt survival of about 13% that is theorized to result through the use of a hatchery. Estimates of the early marine survival are approximate, but sufficient to show a dramatic difference in the response of the two rearing types to the marine nursery area. If the declining trend in production continues for both rearing types, modifications to the hatchery program are needed to improve survival or an emphasis on improving the abundances of wild stocks is necessary, or both. The discovery that the juvenile Cowichan River chinook salmon remain within a relatively confined area of the Gulf Islands within the Strait of Georgia offers an excellent opportunity to research the mechanisms that cause the early marine mortalities and hopefully contribute to a management that improves the production.     

Glochidiosis of salmonid fishes. II. Comparison of tissue response of coho and chinook salmon to experimental infection with Margaritifera margaritifera (L.) (Pelecypoda: Margaritanidae)

Coho salmon (Oncorhynchus kisutch) are more resistant to experimental infection with the glochidia of the freshwater mussel (Margaritifera margaritifera) than are chinook salmon (Oncorhynchus tshawytscha). Histological sections made at intervals during the infection showed that coho salmon sloughed the parasites from their gills by 4.5 days postinfection, but the parasites remained encysted in the gills of chinook salmon for 12 weeks, when metamorphosis to juvenile mussels was complete. Coho salmon sloughed the parasites by a well-developed hyperplasia. No such pronounced hyperplastic reaction was seen in the gills of chinook salmon.     

Effects of handling and electrofishing on plasma glucose and whole blood lactate of Leuciscus cephalus

Chub Leuciscus cephalus exposed to simulated pulsed direct current electrofishing operations exhibited rapid elevations in plasma glucose and blood lactate levels. Plasma glucose levels were significantly higher 0·5 h after simulated electrofishing operations, and peaked 2 h after treatment. Glucose levels remained high for up to 4 h. No changes in plasma glucose were evident following handling. Simulated electrofishing operations and handling induced an immediate lactacidosis in chub. Initial responses to both treatments were similar except that blood lactate was significantly higher in fish exposed to simulated electrofishing operations than in handled fish 5 and 15 min after treatment. Blood lactate remained elevated in fish exposed to simulated electrofishing operations for 2 h, while blood lactate of handled fish returned to levels similar to those in the control fish within 0·5 h post‐treatment.     

Contrasting Functional Performance of Juvenile Salmon Habitat in Recovering Wetlands of the Salmon River Estuary, Oregon, U.S.A.

For an estuarine restoration project to be successful it must reverse anthropogenic effects and restore lost ecosystem functions. Restoration projects that aim to rehabilitate endangered species populations make project success even more important, because if misjudged damage to already weakened populations may result. Determining project success depends on our ability to assess the functional state or “performance” and the trajectory of ecosystem development. Mature system structure is often the desired “end point” of restoration and is assumed to provide maximum benefit for target species; however, few studies have measured linkages between structure and function and possible benefits available from early recovery stages. The Salmon River estuary, Oregon, U.S.A., offers a unique opportunity to simultaneously evaluate several estuarine restoration projects and the response of the marsh community while making comparisons with a concurring undiked portion of the estuary. Dikes installed in three locations in the estuary during the early 1960s were removed in 1978, 1987, and 1996, creating a “space‐for‐time substitution” chronosequence. Analysis of the marsh community responses enables us to use the development state of the three recovering marshes to determine a trajectory of estuarine recovery over 23 years and to make comparisons with a reference marsh. We assessed the rate and pattern of juvenile salmon habitat development in terms of fish density, available prey resources, and diet composition of wild juvenile Oncorhynchus tshawytscha (chinook salmon). Results from the outmigration of 1998 and 1999 show differences in fish densities, prey resources, and diet composition among the four sites. Peaks in chinook salmon densities were greatest in the reference site in 1998 and in the youngest (1996) site in 1999. The 1996 marsh had higher densities of chironomids (insects; average 864/m²) and lower densities of amphipods (crustaceans; average 8/m³) when compared with the other sites. Fauna differences were reflected in the diets of juvenile chinook with those occupying the 1978 and 1996 marshes based on insects (especially chironomids), whereas those from the 1987 and reference marshes were based on crustaceans (especially amphipods). Tracking the development of recovering emergent marsh ecosystems in the Salmon River estuary reveals significant fish and invertebrate response in the first 2 to 3 years after marsh restoration. This pulse of productivity in newly restored systems is part of the trajectory of development and indicates some level of early functionality and the efficacy of restoring estuarine marshes for juvenile salmon habitat. However, to truly know the benefits consumers experience in recovering systems requires further analysis that we will present in forthcoming publications.     

Ecological Risk Assessment Paradigm for Salmon: Analyzing Immune Function to Evaluate Risk

Wild Pacific salmon populations are in serious decline, and as a result, a number of salmon stocks are listed as threatened or endangered under the Endangered Species Act. Our research identifies and supports the possibility that certain environmental contaminants can alter salmon survival, and as a result may contribute to these species being at risk. We have shown that juvenile chinook salmon (Oncorhynchus tshawytscha) are exposed to polychlorinated biphenyls (PCBs) and polycyclic aromatic hydrocarbons (PAHs) as they migrate through a contaminated urban estuary in Puget Sound WA (the Duwamish Waterway estuary). Immune function was analyzed in these fish by examining the ability of their anterior kidney and splenic leukocytes to produce a primary and secondary in vitro plaque-forming cell (PFC) response to the hapten, trinitrophenyl (TNP), and by determining their susceptibility to a marine pathogen, Vibrio anguillarum. We found that fish outmigrating from the urban estuary produced a significantly lower PFC response to TNP and were more susceptible to the pathogen, compared to juvenile salmon collected from a rural estuary during their outmigration. In the laboratory, we exposed juvenile chinook salmon collected from a hatchery to either a PCB technical mixture or a PAH compound to determine if these contaminants have the potential to alter immune function in salmon. Indeed, we found that salmon exposed in the laboratory to either the PCB mixture or the PAH also produced lower PFC responses and were more susceptible to disease compared to animals treated with the solvent vehicle. In summary, contaminants such as PAHs and PCBs are demonstrated to influence salmon health, and thus have the potential to adversely impact salmon populations.     

Behavioral Thermoregulation by Juvenile Spring and Fall Chinook Salmon, Oncorhynchus Tshawytscha, during Smoltification

Fall chinook salmon evolved to emigrate during the summer months. The shift in the temperature preference we observed in smolting fall chinook but not spring chinook salmon may reflect a phylogenetic adaptation to summer emigration by (1) providing directional orientation as fall chinook salmon move into the marine environment, (2) maintaining optimal gill function during emigration and seawater entry, and/or (3) resetting thermoregulatory set-points to support physiological homeostasis once smolted fish enter the marine environment. Phylogenetically determined temperature adaptations and responses to thermal stress may not protect fall chinook salmon from the recent higher summer water temperatures, altered annual thermal regimes, and degraded cold water refugia that result from hydropower regulation of the Columbia and Snake rivers. The long-term survival of fall chinook salmon will likely require restoration of normal annual thermographs and rigorous changes in land use practices to protect critical thermal refugia and control maximum summer water temperatures in reservoirs.     

Juvenile coho salmon track a seasonally shifting thermal mosaic across a river floodplain

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In situ and laboratory studies on the behaviour and survival of Pacific salmon (genus Oncorhynchus)

Juvenile Pacific salmon display a marked surface water orientation during downstream migration, estuarine and nearshore coastal rearing phases. Many estuaries in British Columbia are vertically stratified with a shallow, well-defined halocline which can restrict the dispersion of wastes discharged into less saline surface waters and impose constraints upon aquatic organisms. In situ experiments in an estuary receiving a surface discharge of treated pulp mill wastes, revealed conditions which were lethal to underyearling salmon at, and below the halocline (4.0–6.5 m depth). Behavioural bioassays determined that juvenile chinook salmon were biased towards the water surface and avoided waters at depth. Dissolved oxygen was the variable which affected this distribution most significantly. Surface waters receiving effluent from another pulp mill were lethal to juvenile salmon within 350 m, and a significant vertical avoidance response occurred within 350–950 m of the outfalls. The behavioural response was significantly correlated with in situ temperature, pH and colour (effluent).As a complement to field experiments we developed a 4500 l water column simulator (WCS) to examine salmon behaviour in the laboratory. We investigated the surface water orientation behaviour of juvenile salmon in relation to variations in salinity and dissolved oxygen. Under simulated vertically stratified estuarine conditions, the fish moved freely between overlying fresh water and salt water. Induction of hypoxic conditions in fresh water elicited a downward distribution shift towards the halocline and oxygenated, but more saline, waters. Avoidance reactions (50% level) occurred consistently up to 7–8 mg · l^–1 dissolved oxygen. Salmon continued to examine the hypoxic freshwater zone despite sub-optimal conditions.     

Density‐dependent mortality in Pacific salmon: the ghost of impacts past?

Conservation biologists often ignore density dependence because at‐risk populations are typically small relative to historical levels. However, if populations are reduced as a result of impacts that lower carrying capacity, then density‐dependent mortality may exist at low population abundances. Here, we explore this issue in threatened populations of juvenile chinook salmon (Oncorhynchus tshawytscha). We followed the fate of more than 50 000 juvenile chinook in the Snake River Basin, USA to test the hypothesis that their survival was inversely associated with juvenile density. We also tested the hypotheses that non‐indigenous brook trout and habitat quality affect the presence or strength of density dependence. Our results indicate that juvenile chinook suffer density‐dependent mortality and the strength of density dependence was greater in streams in which brook trout were absent. We were unable to detect an effect of habitat quality on the strength of density dependence. Historical impacts of humans have greatly reduced population sizes of salmon, and the density dependence we report may stem from a shortage of nutrients normally derived from decomposing salmon carcasses. Cohorts of juvenile salmon may experience density‐dependent mortality at population sizes far below historical levels and recovery of imperiled populations may be much slower than currently expected.     

Warming Ocean Conditions Relate to Increased Trophic Requirements of Threatened and Endangered Salmon

The trophic habits, size and condition of yearling Chinook salmon (Oncorhynchus tshawytscha) caught early in their marine residence were examined during 19 survey years (1981–1985; 1998–2011). Juvenile salmon consumed distinct highly piscivorous diets in cold and warm ocean regimes with major differences between ocean regimes driven by changes in consumption of juvenile rockfishes, followed by several other fish prey, adult euphausiids and decapod larvae. Notable, Chinook salmon consumed 30% more food in the warm versus cold ocean regime in both May and June. Additionally, there were about 30% fewer empty stomachs in the warm ocean regime in May, and 10% fewer in warm June periods. The total prey energy density consumed during the warmer ocean regime was also significantly higher than in cold. Chinook salmon had lower condition factor and were smaller in fork length during the warm ocean regime, and were longer and heavier for their size during the cold ocean regime. The significant increase in foraging during the warm ocean regime occurred concurrently with lower available prey biomass. Adult return rates of juvenile Chinook salmon that entered the ocean during a warm ocean regime were lower. Notably, our long term data set contradicts the long held assertion that juvenile salmon eat less in a warm ocean regime when low growth and survival is observed, and when available prey are reduced. Comparing diet changes between decades under variable ocean conditions may assist us in understanding the effects of projected warming ocean regimes on juvenile Chinook salmon and their survival in the ocean environment. Bioenergetically, the salmon appear to require more food resources during warm ocean regimes.     

Prey naivety in the behavioural responses of juvenile Chinook salmon (Oncorhynchus tshawytscha) to an invasive predator

1. Non‐native predators might inflict proportionally higher mortality on prey that have no previous experience of them, compared to species that have coexisted with the predator for some time. 2. We tested whether juvenile Chinook salmon (Oncorhynchus tshawytscha) were less able to recognise a non‐native than a native predator, by investigating behavioural responses to the chemical cues of the invasive smallmouth bass (Micropterus dolomieu) and the native northern pikeminnow (Ptychocheilus oregonensis) in both laboratory and field experiments. 3. Laboratory results demonstrated strong innate antipredator responses of individual juvenile Chinook salmon to northern pikeminnow; fish spent 70% of time motionless and exhibited 100% greater panic response than in controls. By contrast, antipredator responses to the chemical cues of smallmouth bass did not differ from controls. 4. These results were supported by similar differences in recognition of these predator odours by groups of juvenile Chinook salmon in fully natural conditions, though responses reflected a greater range of antipredator behaviours by individuals. In field trials, responses to northern pikeminnow odour resulted in increased flight or absence, reductions in swimming and foraging, and increased time spent near the substratum, compared to smallmouth bass odour. 5. Given that survival of juvenile fish is facilitated by predator recognition, our results support the hypothesis that naivety may be an important factor determining the effect of non‐native predators on prey populations. Efforts to manage the effect of native and non‐native predators may benefit by considering complex behavioural interactions, such as these at the individual and group levels.