Emergence of Baltic salmon, Salmo salar L., in relation to temperature: a laboratory study

Emergence pattern and developmental status at emergence of Baltic salmon fry from the Umeälven hatchery stock (63°50'N, 20°25'E) were studied at 6, 10 and 12°C in the laboratory. The number of days and degree days from hatching to 50% emergence decreased exponentially with increasing temperature. Synchronization of emergence increased with temperature. Optimal temperature for incubation of yolk-sac alevins was 10°C, which resulted in the largest fry at emergence and the lowest death rate. Fry kept at 6°C had the lowest mean weight and at 12°C the highest death rate. The fry emerged at an earlier developmental state with more yolk at 12°C than at 6°C. The Baltic salmon had a faster developmental rate during the gravel-phase, as compared to more southern Atlantic salmon populations.     

Behavioural responses of naive Arctic charr young to chemical cues from salmonid and non-salmonid fish

The ability to distinguish among chemical cues from multiple predators is of key adaptive value for many prey fish. We examined the attractiveness and repulsiveness of chemical stimuli from different coexisting fish species fed on different diets on the behaviour of hatchery reared Arctic charr young in a Y-maze fluviarum, where the charr could choose between two sides either with control water or stimulus water with fish odour. We used stimuli from (1) matching sized conspecifics, large (2) Arctic charr, (3) salmon, (4) brown trout and (5) brown trout fed on Arctic charr fry. Other salmonids were given pellet food. Additional fish odour treatments included piscivorous (6) pike and (7) burbot. In the control trials both sides received control water. Arctic charr young were expected to respond adaptively to the stimuli from coexisting piscivorous fish. The charr most strongly preferred water with the odour of their matching sized conspecifics, which was the only fish odour they were familiar with before the experiments. They also showed significant preference for other salmonid odours, even though these fish are potential predators on small charr. Chemical stimuli from pike and burbot, on the contrary, were strongly avoided, and burbot odour even prevented the charr to swim and enter the lateral halves of the fluviarum. Moreover, odour from brown trout fed on Arctic charr fry was avoided when compared to stimuli from trout fed on pellets. Although the Arctic charr young were completely naive regarding piscivores, the fact that they could distinguish between different predator taxa and diets on the basis of chemical cues only reflects the long coevolutionary history of these fish populations.     

Effects of temperature and a piscivorous fish on diel winter behaviour of juvenile brown trout (Salmo trutta)

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Hatching Time and Alevin Growth Prior to the Onset of Exogenous Feeding in Farmed,Wild and Hybrid Norwegian Atlantic Salmon

The onset of exogenous feeding, when juveniles emerge from the gravel, is a critical event for salmonids where early emergence and large size provide a competitive advantage in the wild. Studying 131 farmed, hybrid and wild Norwegian Atlantic salmon families, originating from four wild populations and two commercial strains, we investigated whether approximately 10 generations of selection for faster growth has also resulted in increased somatic growth prior to the onset of exogenous feeding. In addition, we tested whether relaxed selection in farms has allowed for alterations in hatching time between farmed and wild salmon. Across three cohorts, wild salmon families hatched earlier than farmed salmon families, while hybrid families displayed intermediate hatching times. While the observed differences were small, i.e., 1–15 degree-days (0–3 days, as water temperatures were c. 5–6°C), these data suggest additive genetic variation for hatching time. Alevin length prior to exogenous feeding was positively related to egg size. After removal of egg size effects, no systematic differences in alevin length were observed between the wild and farmed salmon families. While these results indicate additive genetic variation for egg development timing, and wild salmon families consistently hatched earlier than farmed salmon families, these differences were so small they are unlikely to significantly influence early life history competition of farmed and wild salmon in the natural environment. This is especially the case given that the timing of spawning among females can vary by several weeks in some rivers. The general lack of difference in size between farmed and wild alevins, strongly suggest that the documented differences in somatic growth rate between wild and farmed Norwegian Atlantic salmon under hatchery conditions are first detectable after the onset of exogenous feeding.     

Temporal and spatial segregation of spawning in sympatric populations of Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L.

Based on data from Norwegian streams with sympatric populations of Atlantic salmon and brown trout, it is suggested that temporal segregation is the main mechanism segregating Atlantic salmon and brown trout during spawning. Peak spawning of trout was about 15 days earlier than that of salmon. Physical factors, such as water depth, water velocity and distance from the river banks segregate spawning sites of salmon and trout poorly. Gravel sizes of the redds of salmon and trout were significantly different, though with a considerable overlap, and mean egg depth of salmon and trout were 0.18 and 0.12 m, respectively, probably attributable to the different size of spawners of salmon and trout. None of the temporal or spatial parameters analysed segregate spawners of salmon and trout completely. Species determination of eggs and alevins from the redds showed no interspecific superimposition of redds. It is, therefore, concluded that low survival of hybrids after hatching does not explain the low frequency of hybrids observed in sympatric populations of salmon and trout.     

Community structure affects trophic ontogeny in a predatory fish

While most studies have focused on the timing and nature of ontogenetic niche shifts, information is scarce about the effects of community structure on trophic ontogeny of top predators. We investigated how community structure affects ontogenetic niche shifts (i.e., relationships between body length, trophic position, and individual dietary specialization) of a predatory fish, brown trout (Salmo trutta). We used stable isotope and stomach content analyses to test how functional characteristics of lake fish community compositions (competition and prey availability) modulate niche shifts in terms of (i) piscivorous behavior, (ii) trophic position, and (iii) individual dietary specialization. Northern Scandinavian freshwater fish communities were used as a study system, including nine subarctic lakes with contrasting fish community configurations: (i) trout‐only systems, (ii) two‐species systems (brown trout and Arctic charr [Salvelinus alpinus] coexisting), and (iii) three‐species systems (brown trout, Arctic charr, and three‐spined sticklebacks [Gasterosteus aculeatus] coexisting). We expected that the presence of profitable small prey (stickleback) and mixed competitor–prey fish species (charr) supports early piscivory and high individual dietary specialization among trout in multispecies communities, whereas minor ontogenetic shifts were expected in trout‐only systems. From logistic regression models, the presence of a suitable prey fish species (stickleback) emerged as the principal variable determining the size at ontogenetic niche shifts. Generalized additive mixed models indicated that fish community structure shaped ontogenetic niche shifts in trout, with the strongest positive relationships between body length, trophic position, and individual dietary specialization being observed in three‐species communities. Our findings revealed that the presence of a small‐sized prey fish species (stickleback) rather than a mixed competitor–prey fish species (charr) was an important factor affecting the ontogenetic niche‐shift processes of trout. The study demonstrates that community structure may modulate the ontogenetic diet trajectories of and individual niche specialization within a top predator.     

Assessing the reproductive contributions of sympatric anadromous and freshwater‐resident brook trout

The δ¹³C, δ¹⁵N and δ³⁴S isotope values of newly emerged, brook trout Salvelinus fontinalis alevins (free‐swimming individuals beginning exogenous feeding) were examined to determine if progeny of anadromous female spawners could be detected and their contributions to reproduction assessed in river systems with mixed migration strategies. Separation of anadromous and freshwater resident and immigrant sources of progeny could be detected primarily using δ¹³C values before alevins reached a size of 28 mm fork length and c . 20 days after exogenous feeding began. Among 10 populations, the contributions of anadromous females ranged from 0 to 42% of newly emerged alevins. Correlations between anadromous contributions and estimates of total brook trout density and alevin production were unexpectedly negative.     

Ontogenetic niche shifts and resource partitioning in a subarctic piscivore fish guild

The feeding ecology of three piscivorous fish species (perch (Perca fluviatilis), pike (Esox lucius) and burbot (Lota lota)), was studied in the subarctic Pasvik watercourse (69 °N), northern Norway and Russia. All three species primarily occupied the benthic habitats in the watercourse. Perch and burbot exhibited distinct ontogenetic niche shifts in food resource use, perch changing from a dominance of zooplankton to zoobenthos to fish, and burbot from zoobenthos to fish. Fish prey dominated the diet of all the investigated size-classes of pike, but small-sized pike (<20 cm) were not represented in the sample. Fish prey size was positively related to predator size in all three species. Whitefish (Coregonus lavaretus) was the dominant prey of pike and large-sized burbot and perch. Nine-spined sticklebacks (Pungitius pungitius) was also an important prey and appeared to be a dietary stepping-stone enhancing the transition from invertebrate feeding to consumption of large-sized whitefish prey for all three predators. A cluster analysis separated the different size groups of the three predator species into five functional feeding groups, most of them containing two or all three species. Within these feeding groups, and especially among the piscivorous size-classes, there was a strong and significant interspecific overlap in prey selection, and the dietary similarities between the species were in general much larger than the intraspecific similarities between ontogenetic stages. All three piscivorous species are important top predators in the aquatic food web of the watercourse, and their ontogenetic diet shifts and resource partitioning patterns generate a substantial food web complexity in this subarctic ecosystem.     

Responses of tadpoles to hybrid predator odours: strong maternal signatures and the potential risk/response mismatch

Previous studies have established that when a prey animal knows the identity of a particular predator, it can use this knowledge to make an ‘educated guess'' about similar novel predators. Such generalization of predator recognition may be particularly beneficial when prey are exposed to introduced and invasive species of predators or hybrids. Here, we examined generalization of predator recognition for woodfrog tadpoles exposed to novel trout predators. Tadpoles conditioned to recognize tiger trout, a hybrid derived from brown trout and brook trout, showed generalization of recognition of several unknown trout odours. Interestingly, the tadpoles showed stronger responses to odours of brown trout than brook trout. In a second experiment, we found that tadpoles trained to recognize brown trout showed stronger responses to tiger trout than those tadpoles trained to recognize brook trout. Given that tiger trout always have a brown trout mother and a brook trout father, these results suggest a strong maternal signature in trout odours. Tadpoles that were trained to recognize both brown trout and brook trout showed stronger response to novel tiger trout than those trained to recognize only brown trout or only brook trout. This is consistent with a peak shift in recognition, whereby cues that are intermediate between two known cues evoke stronger responses than either known cue. Given that our woodfrog tadpoles have no evolutionary or individual experience with trout, they have no way of knowing whether or not brook trout, brown trout or tiger trout are more dangerous. The differential intensity of responses that we observed to hybrid trout cues and each of the parental species indicates that there is a likely mismatch between risk and anti-predator response intensity. Future work needs to address the critical role of prey naivety on responses to invasive and introduced hybrid predators.     

Arctic charr in sympatry with burbot: ecological and evolutionary consequences

The trophic niche and parasite infection of Arctic charr (Salvelinus alpinus) were explored in two lakes with sympatric burbot (Lota lota) and two lakes without burbot in subarctic Norway. The CPUE of burbot and charr were similar in one lake, but burbot had a low population density in the other. Burbot were benthivorous in both lakes. Other co-occurring species like brown trout (Salmo trutta), Atlantic salmon parr (Salmo salar), grayling (Thymallus thymallus) and minnow (Phoxinus phoxinus) were also benthivores. At high densities, benthivorous burbot forced the whole Arctic charr population to utilise mainly the limnetic trophic niche. In contrast, at low burbot density or without burbot present, Arctic charr were primarily benthivorous in the littoral zone. Thus, a clear interactive segregation in diet was observed between Arctic charr and burbot at high burbot densities. There was also a high predation pressure from burbot on young Arctic charr along the benthic zones. The extensive use of zooplankton as prey caused a high parasite infection pressure of copepod transmitted Diphyllobothrium spp. larvae, with the potential for high negative impact on the Arctic charr population. As the benthivore trophic niche was occupied by burbot, the ecological opportunities for polymorphism with benthivorous ecotypes or morphs of Arctic charr were probably prevented. Therefore, the sympatry with burbot seems to have large ecological and evolutionary consequences for this Arctic charr population compared with neighbouring lakes where burbot is absent.     

The effect of egg size on yolk utilization and growth of rainbow trout alevins (Oncorhynchus mykiss Walbaum)

Changes in body weight and yolk weight were studied in rainbow trout alevins derived from large eggs (diameter >4 mm) and small eggs (egg diameter <4 mm) from the same female. Maximal body weight was reached later, and was higher, for alevins derived from large than from small eggs. The yolk was absorbed more quickly by the alevins of small eggs. Percentage of body water increased in alevins during fasting up to a limit of 91%. Although the limit was the same for both groups of alevins, it was reached more quickly by alevins from small eggs. Relative water content of the yolk did not seem to be influenced by egg size. These results suggest that original egg size had both a quantitative and qualitative effect on the early development of alevins.     

Influence of energetic status on ontogenetic niche shifts: emergence from the redd is linked to metabolic rate in brown trout

Ontogenetic niche shift should occur when the ratio of growth opportunities to mortality risk becomes higher in the subsequent habitat. While most studies have focused on size to understand the timing of these shifts, an endogenous factor like energetic status (interaction between energy available and energy requirements) appears as a natural candidate to integrate and analyze the growth trade-off between habitats. In this study, we measure energetic content and metabolic rate of individual brown trout (Salmo trutta) fry at emergence from gravel to investigate the influence of energetic status on the timing of this critical ontogenetic niche shift. In addition, as offspring energetic status is subject to parental effects, we examine how females could maximize their own fitness by influencing offspring emergence timing. Our results demonstrate that emergence from gravel is influenced by energetic status. Individuals that emerge first have a higher energetic content but deplete it faster because of a higher metabolic rate. We also find that female fecundity is positively related to emergence period duration. Moreover, our results suggest that females may decrease kin competition during the critical period of emergence by influencing the energetic status of offspring, thus, maximizing their own fitness. Our results help elucidate the mechanisms underlying early ontogenetic niche shifts in juvenile fish and suggest reasons why maternal investment can be so variable within populations.     

Water balance trumps ion balance for early marine survival of juvenile pink salmon (Oncorhynchus gorbuscha)

Smolting salmonids typically require weeks to months of physiological preparation in freshwater (FW) before entering seawater (SW). Remarkably, pink salmon (Oncorhynchus gorbuscha) enter SW directly following yolk absorption and gravel emergence at a size of 0.2 g. To survive this exceptional SW migration, pink salmon were hypothesized to develop hypo-osmoregulatory abilities prior to yolk absorption and emergence. To test this, alevins (pre-yolk absorption) and fry (post-yolk absorption) were transferred from FW in darkness to SW under simulated natural photoperiod (SNP). Ionoregulatory status was assessed at 0, 1 and 5 days post-transfer. SW alevins showed no evidence of hypo-osmoregulation, marked by significant water loss and no increase in gill Na?/K?-ATPase (NKA) activity or Na?:K?:2Cl? cotransporter (NKCC) immunoreactive (IR) cell frequency. Conversely, fry maintained water balance, upregulated gill NKA activity by 50 %, increased the NKA α1b/α1a mRNA expression ratio by sixfold and increased NKCC IR cell frequency. We also provide the first evidence of photoperiod-triggered smoltification in pink salmon, as fry exposed to SNP in FW exhibited preparatory changes in gill NKA activity and α1 subunit expression similar to fry exposed to SNP in SW. Interestingly, fry incurred larger increases in whole body Na? than alevins following both SW and FW + SNP exposure (40 and 20 % in fry vs. 0 % in alevins). The ability to incur and tolerate large ion loads may underlie a novel mechanism for maintaining water balance in SW prior to completing hypo-osmoregulatory development. We propose that pink salmon represent a new form of anadromy termed "precocious anadromy".     

Damage released prey alarm substances or predator odours? Risk assessment by an aquatic oligochaete

Although the abilities of prey to detect and respond to chemical substances associated with a predator have been widely reported, the factors promoting the evolution of responses to prey alarm cues vs. predator odours are still vague. In this article, we combined field research with laboratory experiments to explore which chemical substance associated with predator activity (predator odour, conspecific or heterospecific alarm substances) induces defence responses in the aquatic oligochaete Stylaria lacustris, which is vulnerable to common littoral predators. The field results indicated that predators injure the oligochaetes and a great proportion, up to 45% of individuals in the population, were found to be damaged. The results of the laboratory experiments revealed that chemical odours from damselfly larvae feeding on S. lacustris did not induce the defence response in the oligochaetes. On the contrary, oligochaetes detected and responded to alarm substances from damaged conspecifics alone and substances from damaged cladoceran Daphnia magna. We discussed conditions favouring the responses to damage released prey alarm cues instead of predator odours in Stylaria lacustris. Our data suggest that the selection of responses to alarm cues from damaged prey vs. predator odours may be dependent on three factors: (1) non-species-specific predation, (2) divergence of food niche of the different stages of the predator and (3) complex food web with multiple predators. Handling editor: S. Declerk     

Cardiovascular responses of eggs, embryos and alevins of Atlantic salmon and rainbow trout to nitric oxide donors, sodium nitroprusside and isosorbide dinitrate, and inhibitors of nitric oxide synthase, N-nitro-l-arginine methyl ester and aminoguanidine, following short- and long-term exposure

Atlantic salmon embryos and alevins Salmo salar that had been exposed to isosorbide dinitrate (ISDN) for 4 weeks, on transfer to fresh water, showed an increase in heart rate. Unexposed embryos and alevins showed a decrease in heart rate following transfer to 100 μmol l⁻¹ ISDN for 4 h. This is in contrast to adult rainbow trout and higher vertebrates where tachycardia occurred in response to nitric oxide (NO) donors. The decreased heart rate in response to ISDN was inhibited by 2 mg 1⁻¹ methylene blue, indicating that NO activates cardiovascular events via guanylyl cyclase and cyclic guanidine monophosphate. Heart rate of rainbow trout alevins Oncorhynchus mykiss exposed to 100 μmol l⁻¹ aminoguanidine responded with a slowly developed but significant bradycardia over 10 min as did those reared in aminoguanidine for 4 weeks then transferred to fresh water. A potentiated increase in heart rate on exposure to the NO donor sodium nitroprusside (SNP), occurred within 1 min in salmon alevins reared in l -nitro-arginine methyl ester ( l -NAME) for 4 weeks, indicating up-regulation of NO receptors. The evidence for down-regulation of SNP-reared alevins exposed to l -NAME was less well defined. The results suggest that both salmonid embryos and alevins have a functional l -arginine-NO pathway and that NO has a physiological role in control of cardiovascular events.     

Behavioural and Growth Responses of a Territorial Fish (Atlantic Salmon, Salmo salar, L.) to Multiple Predatory Cues

Antipredatory behaviours are important fitness components. The probability of survival decreases if animals fail to respond to signs of danger, but in contrast, energetic costs increase if the response to the threat is exaggerated. We conducted a laboratory experiment designed to examine the behavioural and growth responses of a territorial fish (Atlantic salmon fry, Salmo salar, L.) to different predatory cues (no predatory cues, chemical cues alone, physical cues alone and combined chemical and physical cues). We evaluated the response of Atlantic salmon, focussing on behaviours linked to predator avoidance and to other fitness‐enhancing activities (territory defence and energy acquisition) both during the day and the night. The cost of such responses in terms of growth was assessed and we compared the relative contributions of behaviours in explaining individual growth rate, according to each predation treatment. We demonstrated that the magnitude and nature of behavioural modification varied according to the response variables we considered. An index of predator avoidance and the distance from the food source were affected in an additive fashion by predatory cues (interaction term, p = 0.469 and p = 0.888 for the index of predator avoidance and the distance from the food source respectively); the effect of physical cues was stronger than the effect of chemical cues and the effect of the combined cues was highest. An index of territoriality was affected in a threshold‐like fashion (interaction term, p = 0.040); chemical or physical cues alone had no effect but when both cues were combined, Atlantic salmon significantly reduced their territorial defence. An index of foraging activity was not significantly affected by predatory cues (alone or combined). We detected no effect on the growth rate of Atlantic salmon (p = 0.328). Finally, we found that the relative contribution of behaviours in explaining individual growth rate changed according to the treatments we considered. Overall, these results demonstrated that fish were able to accurately integrate multiple predatory cues and that this information was used to modulate their antipredatory response. Behaviours involved in the response were relatively independent of each other, allowing fish to adopt behavioural tactics that maximized the ratio of net energy gain to predator avoidance.     

Feeding pattern and diet of first feeding brown trout fry under natural conditions

The feeding habits of brown trout Salmo trutta fry were studied during the critical first feeding period in a natural spawning and nursery stream. A low proportion of the fry initiated exogenous feeding before emergence from the gravel, and while having nearly 30% of the yolk sac remaining. This probably reflected low feeding motivation or limited feeding opportunities within the gravel environment. The majority of the fry started feeding after emergence, and after most or all yolk was absorbed. Some fry emerged with large amounts of yolk remaining, while others emerged after yolk exhaustion. The degree of stomach fullness revealed that feeding was more efficient after a territory had been acquired. The diets of the young fry were dominated by chironomid larvae, followed by zooplankton and Plecoptera larvae. Fry dispersing downstream and out of the nursery area were significantly smaller than resident fry, indicating displacement due to competition for territories. The majority of the downstream dispersing fry had initiated feeding, and there was at this point no evidence of starvation in any of the fry. It therefore appeared that the later emerging fry actively migrated out of the overpopulated nursery area to find available territories further downstream.     

Intra- and Interspecific Competition in Hatchery Landlocked Salmon and Brown Trout in Semi-Natural Streams

I studied inter- and intraspecific competition in two hatchery stocks: landlocked salmon with long-hatchery background and a heterogenic brown trout stock. These species are potential competitors in the natural environment when landlocked salmon is being restored to wild by stocking hatchery juveniles. Behavioural responses were studied in four indoor laboratory flumes (400 cm long and 37 cm wide) and habitat use in six semi-natural outdoor streams (26 m long and 1.5 m long). Video recordings were used to monitor fish behaviour and electrofishing for fish positioning in the outdoor channels. The study design included five treatments: two densities of brown trout and salmon in solitary and both species together. The results of the study demonstrated that juvenile brown trout changed their behaviour in laboratory streams in response to presence of the landlocked salmon and the density of the conspecifics also tended to alter the habitat use by brown trout in semi-natural streams. Landlocked salmon juveniles showed no response to treatments. I conclude that possible poor adaptive ability to conditions outside hatchery by the hatchery salmon together and more competitive brown trout stocks may limit the success of management action in restoring landlocked salmon back to their natural streams of stocking.     

Dissolved oxygen concentration and emergence of sea trout fry from natural redds in tributaries of the River Rhine

A total of 4289 sea trout Salmo trutta alevins swam up from six of eleven covered redds. They emerged only from those redds where the mean dissolved oxygen level was >6·9 mg l⁻¹ and the Sorting coe.cient, an indicator of intragravel void space, was <5. These results are discussed in relation to field studies using artificially buried salmonid eggs but also in relation to the ongoing programme for the reintroduction of Atlantic salmon in tributaries of the River Rhine.     

Exposure to a contextually neutral stressor potentiates fear conditioning in juvenile rainbow trout,Oncorhynchus mykiss

Organisms faced with stressors deploy a suite of adaptive responses in the form of behavioral, physiological and cognitive modifications to overcome the challenge. Interactive effects of these responses are known to influence learning and memory processes and facilitation is thought to be dependent, in part, upon contextual relevance of the stressor to the learning task. Predation is one such stressor for prey animals, and their ability to manage reliable information about predators is essential for adaptive antipredator strategies. Here, we investigated (i) the influence cortisol has on the ability of juvenile rainbow trout to learn and retain conditioned antipredator responses to predatory cues, and (ii) whether conditioned behavioral and physiological responses to predator cues are fixed or deployed in a threat-sensitive manner. Trout were fed cortisol-coated pellets minutes prior to a conditioning event where they were exposed to novel predator odor paired with chemical alarm cues (unconditioned stimulus). We tested for conditioned responses by exposing trout to predator cues after 2, 4 or 10 days and subsequently documented physiological and behavioral responses. Both control and cortisol-fed trout learned the predator odor and responded 2 and 4 days post conditioning. However, at 10 days only cortisol-fed trout maintained strong behavioral responses to predator cues. Interestingly, we failed to find conditioned physiological responses to predator odor despite the presence of threat-sensitive cortisol responses to the unconditioned stimulus. Our findings suggest cortisol exposure prior to predator-learning may enhance retention of conditioned responses, even without a contextual link between stressor source and learning task.