Comparing the migration behavior and survival of Atlantic salmon (Salmo salar) and brown trout (Salmo trutta) smolts.

Many organisms rely on migrations between habitats to maximize lifetime fitness, but these migrations can be risky due to a suite of factors. In anadromous salmonids, the smolt migration from fresh water to sea is a critical life stage, during which smolts can experience high mortality from multiple sources. This study investigated the migratory behavior and survival of Atlantic salmon (Salmo salar) and anadromous brown trout (Salmo trutta) smolts during their seaward migration using acoustic telemetry between March and May 2021. Due to the extinction of wild salmon in the River Gudenaa after the construction of the Tange hydropower plant, this study used hatchery-reared salmon originating from a nearby Danish river. A total of 75 hatchery-reared salmon smolts, 75 hatchery-reared trout smolts, and 75 wild trout smolts were tagged with acoustic transmitters and released into River Gudenaa, Denmark. The downstream movements of tagged fish were monitored using acoustic receivers deployed in the river and fjord. Hatchery-reared trout initiated migration first, followed by hatchery-reared salmon, with wild trout being the last to migrate. There was no difference in riverine progression rates among the three smolt groups, but noticeable differences emerged once in the fjord: trout (wild and hatchery) slowed down, whereas hatchery-reared salmon maintained their speed. Riverine migration was predominantly nocturnal for all smolts; however, daytime migration increased at the fjord arrays. Day-of-year significantly influenced diurnal patterns in the river and fjord, where daytime migration increased later in the year. Hatchery-reared salmon and wild trout had reasonably good overall survival from river to sea entry (≥66%), whereas hatchery-reared trout had poor survival (c.26%). The fjord was the major bottleneck for survival of hatchery-reared trout. We found no strong evidence for differences in progression rate or diurnal patterns between wild and hatchery-reared trout to explain the lower survival. This study demonstrates that salmon and trout differ in their life-history strategy already in the post-smolt phase, and that stocking is a sub-optimal strategy to aid wild populations.

Effects of pre-winter cortisol exposure on condition, diet, and morphology of wild juvenile brown trout (Salmo trutta).

Winter is an energetically challenging period for many animals in temperate regions because of the relatively harsh environmental conditions and reduction in food availability during this season. Moreover, stressors experienced by individuals in the fall can affect their subsequent foraging strategy and energy stores after exposure has ended, referred to as carryover effects. We used exogenous cortisol manipulation of wild juvenile brown trout (Salmo trutta) in the fall to simulate a physiological stress response and then investigated short-term (2 weeks) and long-term (4 months) effects on condition metrics (hepatosomatic index and water muscle content), diet (stomach contents and stable isotopes), and morphology during growth in freshwater. We revealed some short-term impacts, likely due to handling stress, and long-term (seasonal) changes in diet, likely reflecting prey availability. Unfortunately, we had very few recaptures of cortisol-treated fish at long-term sampling, limiting detailed analysis about cortisol effects at that time point. Nonetheless, the fish that were sampled showed elevated stable isotopes, suggestive of a cortisol effect long after exposure. This is one of few studies to investigate whether cortisol influences foraging and morphology during juvenile growth, thus extending the knowledge of proximate mechanisms influencing ecologically-relevant phenotypes.

Over 200,000 kilometers of free-flowing river habitat in Europe is altered due to impoundments.

European rivers are disconnected by more than one million man-made barriers that physically limit aquatic species migration and contribute to modification of freshwater habitats. Here, a Conceptual Habitat Alteration Model for Ponding is developed to aid in evaluating the effects of impoundments on fish habitats. Fish communities present in rivers with low human impact and their broad environmental settings enable classification of European rivers into 15 macrohabitat types. These classifications, together with the estimated fish sensitivity to alteration of their habitat are used for assessing the impacts of six main barrier types (dams, weirs, sluices, culverts, fords, and ramps). Our results indicate that over 200,000 km or 10% of previously free-flowing river habitat has been altered due to impoundments. Although they appear less frequently, dams, weirs and sluices cause much more habitat alteration than the other types. Their impact is regionally diverse, which is a function of barrier height, type and density, as well as biogeographical location. This work allows us to foresee what potential environmental gain or loss can be expected with planned barrier management actions in rivers, and to prioritize management actions.

Mechanisms of cardiac collapse at high temperature in a marine teleost (Girella nigrians).

Heat-induced mortality in ectotherms may be attributed to impaired cardiac performance, specifically a collapse in maximum heart rate (fHmax), although the physiological mechanisms driving this phenomenon are still unknown. Here, we tested two proposed factors which may restrict cardiac upper thermal limits: noxious venous blood conditions and oxygen limitation. We hypothesized elevated blood [K+] (hyperkalemia) and low oxygen (hypoxia) would reduce cardiac upper thermal limits in a marine teleost (Girella nigricans), while high oxygen (hyperoxia) would increase thermal limits. We also hypothesized higher acclimation temperatures would exacerbate the harmful effects of an oxygen limitation. Using the Arrhenius breakpoint temperature test, we measured fHmax in acutely warmed fish under control (saline injected) and hyperkalemic conditions (elevated plasma [K+]) while exposed to hyperoxia (200% air saturation), normoxia (100% air saturation), or hypoxia (20% air saturation). We also measured ventricle lactate content and venous blood oxygen partial pressure (PO2) to determine if there were universal thresholds in either metric driving cardiac collapse. Elevated [K+] was not significantly correlated with any cardiac thermal tolerance metric. Hypoxia significantly reduced cardiac upper thermal limits (Arrhenius breakpoint temperature [TAB], peak fHmax, temperature of peak heart rate [TPeak], and temperature at arrhythmia [TARR]). Hyperoxia did not alter cardiac thermal limits compared to normoxia. There was no evidence of a species-wide threshold in ventricular [lactate] or venous PO2. Here, we demonstrate that oxygen limits cardiac thermal tolerance only in instances of hypoxia, but that other physiological processes are responsible for causing temperature-induced heart failure when oxygen is not limited.

The ecological relevance of critical thermal maxima methodology for fishes.

Critical thermal maxima methodology (CTM) has been used to infer acute upper thermal tolerance in fishes since the 1950s, yet its ecological relevance remains debated. In this study, the authors synthesize evidence to identify methodological concerns and common misconceptions that have limited the interpretation of critical thermal maximum (CTmax ; value for an individual fish during one trial) in ecological and evolutionary studies of fishes. They identified limitations of, and opportunities for, using CTmax as a metric in experiments, focusing on rates of thermal ramping, acclimation regimes, thermal safety margins, methodological endpoints, links to performance traits and repeatability. Care must be taken when interpreting CTM in ecological contexts, because the protocol was originally designed for ecotoxicological research with standardized methods to facilitate comparisons within study individuals, across species and contexts. CTM can, however, be used in ecological contexts to predict impacts of environmental warming, but only if parameters influencing thermal limits, such as acclimation temperature or rate of thermal ramping, are taken into account. Applications can include mitigating the effects of climate change, informing infrastructure planning or modelling species distribution, adaptation and/or performance in response to climate-related temperature change. The authors' synthesis points to several key directions for future research that will further aid the application and interpretation of CTM data in ecological contexts.

The Physiological Costs of Reproduction in a Capital Breeding Fish.

AbstractReproduction represents the most energetically demanding period of life for many organisms. Capital breeders, such as anadromous sea trout (Salmo trutta), provide a particularly interesting group of organisms to study within the context of reproduction because they rely on energy stores accrued before breeding to reproduce and sustain all phenotypic and behavioral changes related to reproduction. Energy allocation into current reproduction therefore cannot be mitigated via food intake, resulting in an important life history trade-off. For this reason, exploring indexes related to energetics in salmonids can provide powerful insights into the physiological costs of reproduction. In this study, we sampled blood from and PIT tagged 232 fish captured in the wild before the spawning season. We recaptured and resampled 74 individuals (53 females and 21 males) at the end of the spawning season. Females were further divided into spawning phases (nonspawned, partially spawned, and spawned individuals), though males could not be classified as such. We compared nutritional correlates (triglycerides, cholesterol, calcium, inorganic phosphorus, and total protein), stress correlates (cortisol, sodium, potassium, chloride, and glucose), and indexes of tissue damage (aspartate aminotransferase) between initial capture and recapture as well as among spawning phases in females. We found that nutritional status decreased in all fish throughout the spawning season but that it was substantially lower in females that had spawned. We further found that spawning itself appears stressful, with elevated glucose in partially spawned females and elevated cortisol in male sea trout at recapture. Our findings thus support the idea that the cost of reproduction is energetically high and that incurred stress and a decrease in nutritional status are important physiological costs.

The movement ecology of fishes.

Movement of fishes in the aquatic realm is fundamental to their ecology and survival. Movement can be driven by a variety of biological, physiological and environmental factors occurring across all spatial and temporal scales. The intrinsic capacity of movement to impact fish individually (e.g., foraging) with potential knock-on effects throughout the ecosystem (e.g., food web dynamics) has garnered considerable interest in the field of movement ecology. The advancement of technology in recent decades, in combination with ever-growing threats to freshwater and marine systems, has further spurred empirical research and theoretical considerations. Given the rapid expansion within the field of movement ecology and its significant role in informing management and conservation efforts, a contemporary and multidisciplinary review about the various components influencing movement is outstanding. Using an established conceptual framework for movement ecology as a guide (i.e., Nathan et al., 2008: 19052), we synthesized the environmental and individual factors that affect the movement of fishes. Specifically, internal (e.g., energy acquisition, endocrinology, and homeostasis) and external (biotic and abiotic) environmental elements are discussed, as well as the different processes that influence individual-level (or population) decisions, such as navigation cues, motion capacity, propagation characteristics and group behaviours. In addition to environmental drivers and individual movement factors, we also explored how associated strategies help survival by optimizing physiological and other biological states. Next, we identified how movement ecology is increasingly being incorporated into management and conservation by highlighting the inherent benefits that spatio-temporal fish behaviour imbues into policy, regulatory, and remediation planning. Finally, we considered the future of movement ecology by evaluating ongoing technological innovations and both the challenges and opportunities that these advancements create for scientists and managers. As aquatic ecosystems continue to face alarming climate (and other human-driven) issues that impact animal movements, the comprehensive and multidisciplinary assessment of movement ecology will be instrumental in developing plans to guide research and promote sustainability measures for aquatic resources.

First tagging data on large Atlantic bluefin tuna returning to Nordic waters suggest repeated behaviour and skipped spawning.

Atlantic bluefin tuna (Thunnus thynnus; ABFT) is one of the most iconic fish species in the world. Recently, after being very rare for more than half a century, large bluefin tunas have returned to Nordic waters in late summer and autumn, marking the return of the largest predatory fish in Nordic waters. By tagging 18 bluefin tunas with electronic tags (pop-up satellite archival tags), we show that bluefin tuna observed in Nordic waters undertake different migration routes, with individuals migrating into the western Atlantic Ocean, while others stay exclusively in the eastern Atlantic and enter the Mediterranean Sea to spawn. We additionally present evidence of possible skipped spawning inferred from behavioural analyses. In Nordic waters, ABFT are primarily using the upper water column, likely reflecting feeding activity. The results support the hypothesis that ABFT migrating to Nordic waters return to the same general feeding area within the region on an annual basis. These observations may have important implications for management because (1) tunas that come into Nordic waters might represent only a few year classes (as evidenced by a narrow size range), and thus may be particularly vulnerable to area-specific exploitation, and (2) challenge the assumption of consecutive spawning in adult Atlantic bluefin tuna, as used in current stock assessment models. Without careful management and limited exploitation of this part of the ABFT population, the species' return to Nordic waters could be short-lived.

First tracking of the oceanic spawning migrations of Australasian short-finned eels (Anguilla australis).

Anguillid eel populations have declined dramatically over the last 50 years in many regions of the world, and numerous species are now under threat. A critical life-history phase is migration from freshwater to distant oceans, culminating in a single life-time spawning event. For many anguillids, especially those in the southern hemisphere, mystery still shrouds their oceanic spawning migrations. We investigated the oceanic spawning migrations of the Australasian short-finned eel (Anguilla australis) using pop-up satellite archival tags. Eels were collected from river estuaries (38° S, 142° E) in south-eastern temperate Australia. In 2019, 16 eels were tracked for up to about 5 months, ~ 2620 km from release, and as far north as the tropical Coral Sea (22° S, 155° E) off the north-east coast of Australia. Eels from southern Australia appeared to access deep water off the Australian coast via two main routes: (i) directly east via Bass Strait, or (ii) south-east around Tasmania, which is the shortest route to deep water. Tagged eels exhibited strong diel vertical migrations, alternating between the warm euphotic zone (~ 100-300 m, 15-20 °C) at night and the mesopelagic zone (~ 700-900 m, 6-8 °C) during the day. Marine predators, probably lamnid sharks, tuna, or marine mammals, ended many eel migrations (at least ~ 30%), largely before the eels had left the Australian continental shelf. The long and risky marine migrations of Australasian eels highlight the need for better information on the processes contributing to eel mortality throughout the life cycle, including the impacts of future changes to oceanic currents, predator abundance and direct anthropogenic disturbances.

No Evidence for Long-Term Carryover Effects in a Wild Salmonid Fish.

AbstractEarly-life experiences can shape life histories and population dynamics of wild animals. To examine whether stressful stimuli experienced in early life resulted in carryover effects in later life stages, we conducted several experimental manipulations and then monitored wild fish with passive integrated transponder tags during juvenile out-migration and adult return migration. In total, 3,217 juvenile brown trout (Salmo trutta) were subjected to one of six manipulations: chase to exhaustion, thermal challenge, food deprivation, low-concentration cortisol injection, high-concentration cortisol injection, and sham injection, plus a control group. Cortisol and food deprivation treatments were previously shown to have short-term effects on juveniles, such as lower survival to out-migration and changes in migration timing. However, it remained unknown whether any of the six manipulations had effects that carried over into the adult phase. We therefore investigated whether these extrinsic manipulations, as well as intrinsic factors (size and condition), affected probability of return as adults and time spent at sea. Of the 1,273 fish that out-migrated, 146 returned as adults. We failed to detect any effect of treatments on return rates, while high-concentration cortisol weakly affected time spent at sea in one tagging event. We also found that juvenile condition was positively correlated to likelihood of adult return in only one tagging event. Overall, our findings did not identify either intrinsic factors or extrinsic stressful early-life experiences that have strong effects on fish that survive to adulthood. This suggests that some species may be more resilient than others to stressful stimuli encountered early in life.

Life-history strategies in salmonids: the role of physiology and its consequences.

Salmonids are some of the most widely studied species of fish worldwide. They span freshwater rivers and lakes to fjords and oceans; they include short- and long-distance anadromous migrants, as well as partially migratory and non-migratory populations; and exhibit both semelparous and iteroparous reproduction. Salmonid life-history strategies represent some of the most diverse on the planet. For this reason, salmonids provide an especially interesting model to study the drivers of these different life-history pathways. Over the past few decades, numerous studies and reviews have been published, although most have focused on ultimate considerations where expected reproductive success of different developmental or life-history strategies are compared. Those that considered proximate causes generally focused on genetics or the environment, with less consideration of physiology. Our objective was therefore to review the existing literature on the role of physiology as a proximate driver for life-history strategies in salmonids. This link is necessary to explore since physiology is at the core of biological processes influencing energy acquisition and allocation. Energy acquisition and allocation processes, in turn, can affect life histories. We find that life-history strategies are driven by a range of physiological processes, ranging from metabolism and nutritional status to endocrinology. Our review revealed that the role of these physiological processes can vary across species and individuals depending on the life-history decision(s) to be made. In addition, while findings sometimes vary by species, results appear to be consistent in species with similar life cycles. We conclude that despite much work having been conducted on the topic, the study of physiology and its role in determining life-history strategies in salmonids remains somewhat unexplored, particularly for char and trout (excluding brown trout) species. Understanding these mechanistic links is necessary if we are to understand adequately how changing environments will impact salmonid populations.

More than one million barriers fragment Europe's rivers.

Rivers support some of Earth's richest biodiversity1 and provide essential ecosystem services to society2, but they are often fragmented by barriers to free flow3. In Europe, attempts to quantify river connectivity have been hampered by the absence of a harmonized barrier database. Here we show that there are at least 1.2 million instream barriers in 36 European countries (with a mean density of 0.74 barriers per kilometre), 68 per cent of which are structures less than two metres in height that are often overlooked. Standardized walkover surveys along 2,715 kilometres of stream length for 147 rivers indicate that existing records underestimate barrier numbers by about 61 per cent. The highest barrier densities occur in the heavily modified rivers of central Europe and the lowest barrier densities occur in the most remote, sparsely populated alpine areas. Across Europe, the main predictors of barrier density are agricultural pressure, density of river-road crossings, extent of surface water and elevation. Relatively unfragmented rivers are still found in the Balkans, the Baltic states and parts of Scandinavia and southern Europe, but these require urgent protection from proposed dam developments. Our findings could inform the implementation of the EU Biodiversity Strategy, which aims to reconnect 25,000 kilometres of Europe's rivers by 2030, but achieving this will require a paradigm shift in river restoration that recognizes the widespread impacts caused by small barriers.

The Value of Experimental Approaches in Migration Biology.

The past several decades have ushered in a golden age in the study of migration biology, leading to a wealth of descriptive articles that characterize various aspects of migration and its implications for individuals, populations, and ecosystems. However, relatively few studies have adopted an experimental approach to the study of migration, and fewer still have combined lab and field experiments to glean insights into the mechanisms underlying variation in migration behavior and success. Understanding the proximate and ultimate causes of migration timing, energy allocation and optimization, migration success, and fitness is important to aid the conservation and management of wildlife populations by establishing appropriate protections or managing environmental conditions that influence migration. With recent technological advances and miniaturization of animal-borne electronic tracking devices, as well as ground-, water-, and space-based telemetry infrastructure, researchers have the tools necessary to experimentally test hypotheses central to the mechanics of migrations and individual variation therein. By pairing physiological measurements, molecular analyses, and other approaches within an experimental framework, there is the potential to understand not only how animal migrations function but also what differentiates successful migrations from failed migrations and the associated fitness implications. Experimental approaches to migration biology are particularly important, as they will help us to better comprehend and hopefully predict animal responses to environmental and anthropogenic changes by isolating confounding variables that challenge inferences from observations.

Catchment-scale effects of river fragmentation: A case study on restoring connectivity.

More than two thirds of large rivers worldwide are fragmented, threatening freshwater biodiversity, river integrity, and the services that freshwater ecosystems provide for human populations around the globe. In an effort to alleviate the impacts of barriers, engineered solutions have been developed, though with somewhat underwhelming results. River restoration, especially dam removal, is viewed as the optimal option though seldom the go-to approach. In this study, we evaluated the effects of a large restoration project (pseudo dam removal) in River Kolding, Southern Jutland, Denmark, via a before-after-control-impact (BACI) approach. Using a large dataset of electrofishing data from 74 sites (including downstream unaffected sites, reconnected sites and upstream regulated sites), we found that habitat connectivity was restored successfully, with a large increase in young-of-the-year brown trout (Salmo trutta) at reconnected sites, reaching similar densities to downstream (non-affected) sites. We further observed a decrease in length at reconnected sites, suggesting that natural spawning and rearing habitats were successfully restored too.

Cortisol predicts migration timing and success in both Atlantic salmon and sea trout kelts.

Kelts - individuals of anadromous fish species which have successfully spawned and may return to sea to repeat the cycle - are perhaps the least studied life stage of iteroparous fish species. To date, our understanding of what makes them successful in their return migration to sea is limited. We investigated the relationship between three physiological parameters (baseline cortisol, baseline glucose and low molecular weight antioxidants) and the timing and success of Atlantic salmon (Salmo salar) and sea trout (Salmo trutta) kelt migration. To do so, we combined blood samples obtained within 3 minutes of capture and acoustic telemetry to track 66 salmon and 72 sea trout as they migrated out of rivers, into fjords and out at sea. We show that baseline cortisol may be a good predictor of migration success. Individuals with high baseline cortisol levels exited the river earlier but were less likely to successfully reach the sea. Similar relationships were not observed with glucose or antioxidants. We provide the first evidence to support the role of physiological status in migration success in Atlantic salmon and sea trout kelts. Our findings contribute to our understanding of the relationship between physiology and fitness in wild animals. Further, we suggest that migration timing is a trade-off between stress and readiness to migrate.

N-acetylcysteine manipulation fails to elicit an increase in glutathione in a teleost model.

Levels of oxidative stress can be affected by a range of compounds including toxins and pharmaceuticals. Antioxidants are important protective compounds which counteract the damaging effects of oxidative stress. Glutathione (GSH) is one of the main antioxidants for many organisms and can be synthesized from administered N-acetylcysteine (NAC). NAC has therefore often been used in a wide range of taxa to manipulate levels of GSH. Our objective was to validate this approach in a wild temperate teleost fish model, the brown trout (Salmo trutta). We used intracoelomic injections of NAC in saline and vegetable shortening, at two different concentrations (100 and 400 mg/kg), with the appropriate controls and shams, under controlled laboratory settings. We found that NAC failed to elicit an increase in GSH over three time periods and concluded that NAC is not an effective method to enhance GSH levels in teleost fish using the concentrations and vehicles tested here. We emphasize the importance of validation studies across all new species/taxa when possible and suggest that more investigation is required with regard to NAC manipulation in fish if this approach is to be used.