Association of ectoparasite Lepeophtheirus salmonis counts on farmed Atlantic salmon and wild sea trout in Scotland.

Parasitic sea lice (Copepoda: Caligidae) colonising marine salmonid (Salmoniformes: Salmonidae) aquaculture production facilities have been implicated as a possible pressure on wild salmon and sea trout populations. This investigation uses monitoring data from the mainland west coast and Western Isles of Scotland to estimate the association of the abundance of adult female Lepeophtheirus salmonis (Krøyer) colonising farmed Atlantic salmon Salmo salar L. with the occurrence of juvenile and mobile L. salmonis on wild sea trout, anadromous S. trutta L. The associations were evaluated using generalised linear mixed models incorporating farmed adult female salmon louse abundances which are temporally lagged relative to dependent wild trout values. The pattern of lags, which is consistent with time for L. salmonis development between egg and infective stage, was evaluated using model deviances. A significant positive association is identified between adult female L. salmonis abundance on farms and juvenile L. salmonis on wild trout. This association is consistent with a causal relationship in which increases in the number of L. salmonis copepodids originating from lice colonising farmed Atlantic salmon cause an increase of L. salmonis abundance on wild sea trout.

High summer temperatures are associated with poorer performance of underyearling Atlantic salmon (Salmo salar) in upland streams.

Future warming scenarios are predicted to result in an increased frequency of high, and potentially stressful, temperatures in aquatic ecosystems. Here we examined whether the performance of wild underyearling Atlantic salmon (Salmo salar) in Scottish streams stocked with identical egg densities was influenced by thermal stress. Biomass and density declined with degree hours exceeding 23°C, indicating apparent mortality or emigration as a possible result of exposure to high temperatures. These results strengthen the need for further action such as riparian tree planting to reduce stream summer temperatures.

Nutrient limitation in Atlantic salmon rivers and streams: Causes, consequences, and management strategies.

Freshwater catchments can experience nutrient deficits that result in reduced primary and secondary productivity. The most commonly limiting nutrients are nitrogen and phosphorus, either separately or together. This review considers the impact of increasing nutrient limitation in temperate basin stream and river systems, focusing on upland areas that currently or previously supported wild Atlantic salmon (Salmo salar) populations.Anthropogenic changes to land use and increases in river barriers have altered upland nutrient dynamics, with particular impacts on salmon and other migratory fish species which may be net importers of nutrients to upland streams. Declining salmon populations may further reduce nutrient sources, reducing ecosystem and fisheries productivity below desired levels.Experimental manipulations of nutrient levels have examined the impacts of this cultural oligotrophication. There is evidence that growth and biomass of juvenile salmon can be increased via appropriate additions of nutrients, offering potential as a conservation tool. However, further research is required to understand the long-term effects of these additions on salmon populations and stream ecosystems, and to assess the vulnerability of downstream habitats to eutrophication as a result.Although purposeful nutrient addition with the aim of enhancing and conserving salmonid populations may be justified in some cases, it should be undertaken in an adaptive management framework. In addition, nutrient addition should be linked to nutrient retention and processing, and integrated into large-scale habitat restoration and recovery efforts.Both the scientific and the management community should recognize that the ecological costs and benefits associated with adding nutrients to salmon streams may change in a non-stationary world.

Metabolic Rate Interacts with Resource Availability to Determine Individual Variation in Microhabitat Use in the Wild.

Ecological pressures such as competition can lead individuals within a population to partition resources or habitats, but the underlying intrinsic mechanisms that determine an individual's resource use are not well understood. Here we show that an individual's own energy demand and associated competitive ability influence its resource use, but only when food is more limiting. We tested whether intraspecific variation in metabolic rate leads to microhabitat partitioning among juvenile Atlantic salmon (Salmo salar) in natural streams subjected to manipulated nutrient levels and subsequent per capita food availability. We found that individual salmon from families with a higher baseline (standard) metabolic rate (which is associated with greater competitive ability) tended to occupy faster-flowing water, but only in streams with lower per capita food availability. Faster-flowing microhabitats yield more food, but high metabolic rate fish only benefited from faster growth in streams with high food levels, presumably because in low-food environments the cost of a high metabolism offsets the benefits of acquiring a productive microhabitat. The benefits of a given metabolic rate were thus context dependent. These results demonstrate that intraspecific variation in metabolic rate can interact with resource availability to determine the spatial structuring of wild populations.

Adaptive Maternal Investment in the Wild? Links between Maternal Growth Trajectory and Offspring Size, Growth, and Survival in Contrasting Environments.

Life-history theory predicts that investment per offspring should correlate negatively with the quality of the environment that offspring are anticipated to encounter; parents may use their own experience as juveniles to predict this environment and may modulate offspring traits, such as growth capacity and initial size. We manipulated nutrient levels in the juvenile habitat of wild Atlantic salmon (Salmo salar) to investigate the hypothesis that the egg size that maximizes juvenile growth and survival depends on environmental quality. We also tested whether offspring traits were related to parental growth trajectory. Mothers that grew fast when young produced more offspring and smaller offspring than mothers that grew slowly to reach the same size. Despite their size disadvantage, offspring of faster-growing mothers grew faster than those of slower-growing mothers in all environments, counter to the expectation that they would be competitively disadvantaged. However, they had lower relative survival in environments where the density of older predatory/competitor fish was relatively high. These links between maternal (but not paternal) growth trajectory and offspring survival rate were independent of egg size, underscoring that mothers may be adjusting egg traits other than size to suit the environment their offspring are anticipated to face.

Effective methodologies for enantiomeric separations of 150 pharmacology and toxicology related 1°, 2°, and 3° amines with core-shell chiral stationary phases.

Core-shell particles (superficially porous particles, SPPs) have been proven to provide high-throughput and effective separations of a variety of chiral molecules. However, due to their limited commercialization, many separations have not been reported with these stationary phases. In this study, four SPP chiral stationary phases (CSPs) were utilized for the enantiomeric separation of 150 chiral amines. These amines encompass a variety of structural and drug classes, which are particularly important to the pharmaceutical industry and in forensics. This comprehensive evaluation demonstrates the power of these CSPs and the ease of method development and optimization. The CSPs used in this study included the macrocyclic glycopeptide-based CSPs (VancoShell and NicoShell), the cyclodextrin-based CSP (CDShell-RSP), and the cyclofructan-based CSP (LarihcShell-P). These CSPs offered versatility for a variety of applications and worked in a complementary fashion to baseline separate all 150 amines. The LarihcShell-P was highly effective for separating primary amines. VancoShell, NicoShell, and CDShell-RSP were useful for separating all types of amines. These CSPs are multi-modal and can be utilized with mass spectrometry compatible solvents. Eighteen racemic controlled substances were simultaneously baseline separated in a single liquid chromatography-mass spectrometry (LC-MS) analysis. Details in high-performance liquid chromatography (HPLC) parameters will be discussed as well as the improved chromatographic performance afforded by the SPP CSPs.

Telomere elongation during early development is independent of environmental temperatures in Atlantic salmon.

There is increasing evidence from endothermic vertebrates that telomeres, which cap the ends of chromosomes and play an important role in chromosome protection, decline in length during postnatal life and are a useful indicator of physiological state and expected lifespan. However, much less is currently known about telomere dynamics in ectothermic vertebrates, which are likely to differ from that of endotherms, at least in part due to the sensitivity of ectotherm physiology to environmental temperature. We report here on an experiment in which Atlantic salmon (Salmo salar) were reared through the embryonic and larval stages of development, and under differing temperatures, in order to examine the effects of environmental temperature during early life on telomere dynamics, oxidative DNA damage and cellular proliferation. Telomere length significantly increased between the embryonic and larval stages of development. Contrary to our expectations, variation in telomere length at the end of the larval stage was unrelated to either cell proliferation rate or the relative level of oxidative DNA damage, and did not vary between the temperature treatments. This study suggests that salmon are able to restore the length of their telomeres during early development, which may possibly help to buffer potentially harmful environmental effects experienced in early life.

Links between parental life histories of wild salmon and the telomere lengths of their offspring.

The importance of parental contributions to offspring development and subsequent performance is self-evident at a genomic level; however, parents can also affect offspring fitness by indirect genetic and environmental routes. The life history strategy that an individual adopts will be influenced by both genes and environment; and this may have important consequences for offspring. Recent research has linked telomere dynamics (i.e., telomere length and loss) in early life to future viability and longevity. Moreover, a number of studies have reported a heritable component to telomere length across a range of vertebrates, although the effects of other parental contribution pathways have been far less studied. Using wild Atlantic salmon with different parental life histories in an experimental split-brood in vitro fertilization mating design and rearing the resulting families under standardized conditions, we show that there can be significant links between parental life history and offspring telomere length (studied at the embryo and fry stage). Maternal life history traits, in particular egg size, were most strongly related to offspring telomere length at the embryonic stage, but then became weaker through development. In contrast, paternal life history traits, such as the father's growth rate in early life, had a greater association in the later stages of offspring development. However, offspring telomere length was not significantly related to either maternal or paternal age at reproduction, nor to paternal sperm telomere length. This study demonstrates both the complexity and the importance of parental factors that can influence telomere length in early life.

Nutrients from salmon parents alter selection pressures on their offspring.

Organisms can modify their surrounding environment, but whether these changes are large enough to feed back and alter their evolutionary trajectories is not well understood, particularly in wild populations. Here we show that nutrient pulses from decomposing Atlantic salmon (Salmo salar) parents alter selection pressures on their offspring with important consequences for their phenotypic and genetic diversity. We found a strong survival advantage to larger eggs and faster juvenile metabolic rates in streams lacking carcasses but not in streams containing this parental nutrient input. Differences in selection intensities led to significant phenotypic divergence in these two traits among stream types. Stronger selection in streams with low parental nutrient input also decreased the number of surviving families compared to streams with high parental nutrient levels. Observed effects of parent-derived nutrients on selection pressures provide experimental evidence for key components of eco-evolutionary feedbacks in wild populations.

Maternal age at maturation underpins contrasting behavior in offspring.

In species where parental care occurs primarily via the provisioning of eggs, older females tend to produce larger offspring that have better fitness prospects. Remarkably however, a relationship between age of mother and fitness of offspring has also been reported independently of effects on offspring size suggesting that there may be other factors at play. Here, using experimental matings between wild Atlantic salmon that differed in their age at sexual maturation, we demonstrate distinct size-independent variation in the behavior of their offspring that was related to the maturation age of the mother (but not the father). We found that when juvenile salmon were competing for feeding territories, offspring of early-maturing mothers were more aggressive than those of late-maturing mothers, but were out-competed for food by them. This is the first demonstration of a link between natural variation in parental age at maturation and variation in offspring behavior.

Basin-scale phenology and effects of climate variability on global timing of initial seaward migration of Atlantic salmon (Salmo salar).

Migrations between different habitats are key events in the lives of many organisms. Such movements involve annually recurring travel over long distances usually triggered by seasonal changes in the environment. Often, the migration is associated with travel to or from reproduction areas to regions of growth. Young anadromous Atlantic salmon (Salmo salar) emigrate from freshwater nursery areas during spring and early summer to feed and grow in the North Atlantic Ocean. The transition from the freshwater ('parr') stage to the migratory stage where they descend streams and enter salt water ('smolt') is characterized by morphological, physiological and behavioural changes where the timing of this parr-smolt transition is cued by photoperiod and water temperature. Environmental conditions in the freshwater habitat control the downstream migration and contribute to within- and among-river variation in migratory timing. Moreover, the timing of the freshwater emigration has likely evolved to meet environmental conditions in the ocean as these affect growth and survival of the post-smolts. Using generalized additive mixed-effects modelling, we analysed spatio-temporal variations in the dates of downstream smolt migration in 67 rivers throughout the North Atlantic during the last five decades and found that migrations were earlier in populations in the east than the west. After accounting for this spatial effect, the initiation of the downstream migration among rivers was positively associated with freshwater temperatures, up to about 10 °C and levelling off at higher values, and with sea-surface temperatures. Earlier migration occurred when river discharge levels were low but increasing. On average, the initiation of the smolt seaward migration has occurred 2.5 days earlier per decade throughout the basin of the North Atlantic. This shift in phenology matches changes in air, river, and ocean temperatures, suggesting that Atlantic salmon emigration is responding to the current global climate changes.

Spatial variation in the relationship between performance and metabolic rate in wild juvenile Atlantic salmon.

Maintenance of metabolic rate (MR, the energy cost of self-maintenance) is linked to behavioural traits and fitness and varies substantially within populations. Despite having received much attention, the causes and consequences of this variation remain obscure. Theoretically, such within-population variation in fitness-related traits can be maintained by environmental heterogeneity in selection patterns, but for MR, this has rarely been tested in nature. Here, we experimentally test whether the relationship between MR and performance can vary spatially by assessing survival, growth rate and movement of Atlantic salmon (Salmo salar L.) juveniles from 10 family groups differing in MR (measured as egg metabolism) that were stocked in parallel across 10 tributaries of a single watershed. The relationship between MR and relative survival and growth rate varied significantly among tributaries. Specifically, the effect of MR ranged from negative to positive for relative survival, whereas it was negative for growth rate. The association between MR and movement was positive and did not vary significantly among tributaries. These results are consistent with a fitness cost of traits associated with behavioural dominance that varies across relatively small spatial scales (within a single watershed). More generally, our results support the hypothesis that spatial heterogeneity in environmental conditions contributes to maintain within-population variation in fitness-related traits, such as MR.

The performance advantage of a high resting metabolic rate in juvenile salmon is habitat dependent.

1. Basal levels of metabolism vary significantly among individuals in many taxa, but the effects of this on fitness are generally unknown. Resting metabolic rate (RMR) in juvenile salmon and trout is positively related to dominance status and ability to obtain a feeding territory, but it is not clear how this translates into performance in natural conditions. 2. The relationships between RMR, dominance, territoriality and growth rates of yearling Atlantic salmon Salmo salar were examined in relation to predictability in food supply and habitat complexity, using replicate sections of a large-scale controlled semi-natural stream. 3. Estimated RMR was a strong predictor of dominance, and under conditions of a predictable food supply in a structurally simple habitat, high estimated RMR fish obtained the best feeding territories and grew faster. 4. When the spatial distribution of food was made less predictable, dominant (high estimated RMR) fish were still able to occupy the most profitable feeding locations by periodically moving location to track the changes in food availability, but RMR was no longer a predictor of growth rate. Moreover, when a less predictable food supply was combined with a visually more complex (and realistic) habitat, fish were unable to track changes in food availability, grew more slowly and exhibited greater site fidelity, and there were no relationships between estimated RMR and quality of occupied territory or growth rate. 5. The relative benefit of RMR is thus context dependent, depending on both habitat complexity and the predictability of the food supply. Higher habitat complexity and lower food predictability decrease the performance advantages associated with a high RMR.

The spatial scale of competition from recruits on an older cohort in Atlantic salmon.

Competitive effects of younger cohorts on older ones are frequently assumed to be negligible in species where older, larger individuals dominate in pairwise behavioural interactions. Here, we provide field estimates of such competition by recruits on an older age class in Atlantic salmon (Salmo salar), a species where observational studies have documented strong body size advantages which should favour older individuals in direct interactions. By creating realistic levels of spatial variation in the density of underyearling (YOY) recruits over a 1-km stretch of a stream, and obtaining accurate measurements of individual growth rates of overyearlings (parr) from capture-mark-recapture data on a fine spatial scale, we demonstrate that high YOY density can substantially decrease parr growth. Models integrating multiple spatial scales indicated that parr were influenced by YOY density within 16 m. The preferred model suggested parr daily mass increase to be reduced by 39% when increasing YOY density from 0.0 to 1.0 m(-2), which is well within the range of naturally occurring densities. Reduced juvenile growth rates will in general be expected to reduce juvenile survival (via increased length of exposure to freshwater mortality) and increase generation times (via increased age at seaward migrations). Thus, increased recruitment can significantly affect the performance of older cohorts, with important implications for population dynamics. Our results highlight that, even for the wide range of organisms that rely on defendable resources, the direction of competition among age classes cannot be assumed a priori or be inferred from behavioural observations alone.

The spatial scale of density-dependent growth and implications for dispersal from nests in juvenile Atlantic salmon.

By dispersing from localized aggregations of recruits, individuals may obtain energetic benefits due to reduced experienced density. However, this will depend on the spatial scale over which individuals compete. Here, we quantify this scale for juvenile Atlantic salmon (Salmo salar) following emergence and dispersal from nests. A single nest was placed in each of ten replicate streams during winter, and information on the individual positions (±1 m) and the body sizes of the resulting young-of-the-year (YOY) juveniles was obtained by sampling during the summer. In six of the ten streams, model comparisons suggested that individual body size was most closely related to the density within a mean distance of 11 m (range 2-26 m). A link between body size and density on such a restricted spatial scale suggests that dispersal from nests confers energetic benefits that can counterbalance any survival costs. For the four remaining streams, which had a high abundance of trout and older salmon cohorts, no single spatial scale could best describe the relation between YOY density and body size. Energetic benefits of dispersal associated with reduced local density therefore appear to depend on the abundance of competing cohorts or species, which have spatial distributions that are less predictable in terms of distance from nests. Thus, given a trade-off between costs and benefits associated with dispersal, and variation in benefits among environments, we predict an evolving and/or phenotypically plastic growth rate threshold which determines when an individual decides to disperse from areas of high local density.

Divergent trends in anadromous salmonid populations in Norwegian and Scottish rivers.

The Atlantic salmon (Salmo salar) is a charismatic anadromous fish of high conservation and economic value. Concerns have been expressed regarding the long-term viability of fisheries throughout the species's distributional range because of abundance variations that cannot currently be explained or predicted. Here, we analyse long-term catch data obtained over a wide geographical range and across a range of spatial subscales to understand more fully the factors that drive population abundance. We use rod catch data from 84 Norwegian rivers over 125 years (1876-2000) and 48 Scottish rivers over 51 years (1952-2002). The temporal correlation in catches is very long-term, with trends persisting over several decades. The spatial correlation is relatively short-range, indicating strong local-scale effects on catch. Furthermore, Scottish salmon populations exhibit recent negative trends in contrast to some more positive trends in Norway--especially in the north.

Nest distribution shaping within-stream variation in Atlantic salmon juvenile abundance and competition over small spatial scales.

1. Spatial heterogeneity in population density is predicted to have important effects on population characteristics, such as competition intensity and carrying capacity. Patchy breeding distributions will tend to increase spatial heterogeneity in population density, whereas dispersal from breeding patches will tend to decrease it. The potential for dispersal to homogenize densities is likely to differ both among organisms (e.g. plants vs. mobile animals) and throughout ontogeny (e.g. larvae vs. adults). However, for mobile organisms, experimental studies of the importance of breeding distributions from the wild are largely lacking. 2. In the present study, experimental manipulations replicated over eight natural streams and 2 years enabled us to test for effects of the distribution of Atlantic salmon eggs over spatial scales which are relevant to local interactions among individuals. Artificial nests were placed along 250 m study reaches at one of two levels of nest dispersion - patchy (two nests per stream) and dispersed (10 nests per stream) - while holding total egg density (eggs m(-2) stream area) constant. 3. Nest dispersion had significant effects on the spatial distribution of the resulting juveniles in their first summer. Patchy nest distributions resulted in a highly right-skewed frequency distribution of local under-yearling densities (among 25 m sampling sections), as sample sections adjacent to the nest sites had relatively high densities. In contrast, dispersed nest distributions yielded approximately normal density distributions. Sections with high relative densities in the patchy nest distribution treatments also had relatively small juvenile body sizes, and patchy egg distribution appeared to produce a higher redistribution of individuals from the first to the second juvenile growth season than the dispersed distribution. 4. Because patchy breeding distribution combined with limited early dispersal can create spatial variation in density over scales directly relevant for individual interactions, this will be one important component in determining mean levels of early juvenile competition and its spatial variation within populations. Assuming random or ideal-free distribution of individuals may therefore underestimate the mean level of density experienced by juveniles over surprisingly small spatial scales (orders of magnitude smaller than total spatial extent of populations), even for mobile organisms.

Effects of food availability on temporal activity patterns and growth of Atlantic salmon.

1. Patterns of sheltering and activity are of fundamental importance in the ecology of animals and in determining interactions among predators and prey. Balancing decreased mortality risk when sheltering with increased feeding rate when exposed is believed to be a key determinant of diel patterns of sheltering in many animals. 2. Despite lower foraging efficiency at night than during the day, Atlantic salmon Salmo salar parr are nocturnal during winter and at low summer temperatures. Nocturnal activity also occurs at warm water temperatures during summer, but little is known about the functional significance of this behaviour. 3. This study aimed to determine: (1) the preferred activity and shelter pattern of Atlantic salmon parr during warm summer months, and (2) their response to variations in food availability when balancing growth rate (G) and mortality risk (M), as expressed through time out of shelter. We differentiated among four potential responses to reduced food availability: (1) no response; (2) G decreases but M remains constant; (3) G remains constant but M increases; and (4) G decreases and M increases. 4. Time exposed from shelter was inversely related to food availability. Fish subject to high food availability were significantly less active during the day than those with restricted rations. However, food availability had no significant effect on the extent to which fish were active at night. There was no evidence of variation in growth rate with food availability. 5. Salmon were predominantly nocturnal at high ration levels, consistent with their previously reported behaviour during winter. Rather than switching to diurnal behaviour at high temperatures per se, as previously was supposed, it appears that the fish are diurnal only to the extent needed to sustain a growth rate, and this extent depends on food availability. 6. Atlantic salmon parr modulate the amount of time they are active rather than growth when responding to variations in food availability over an order of magnitude.