Changes of sperm morphology, volume, density, and motility parameters in northern pike during the spawning period.

Sexually mature males (BW = 1600 ± 150 g and TL = 235 ± 30 mm) of northern pike (Esox lucius L.) were randomly selected from a pond to record changes in their sperm quality parameters (spermatozoa morphology, sperm volume, density, and motility parameters) during the spawning season. The morphological and motility parameters changed significantly during the reproductive season with following trends. Only, head width was not changed during the spawning season. The longest spermatozoa and its flagellar length were found at the middle of spawning period (TL = 38.24 ± 0.37 µm and 35.14 ± 0.26 µm) and shortest at the beginning of spawning period (TL = 34.81 ± 0.29 µm and 32.53 ± 0.18 µm). Other morphological characters were always the lowest at the beginning of spawning period. Sperm volume was changed from 0.33 ± 0.3 ml in February, 0.43 ± 0.2 ml in March to 0.24 ± 0.1 ml in April, and density from 16.2 ± 0.2 × 109 spermatozoa ml-1 in February, 19.4 ± 0.2 × 109 spermatozoa ml-1 in March to 4.8 ± 0.2 × 109 spermatozoa ml-1 in April. Same sperm velocity was observed in all spawning terms at 10 and 20 s after activation. Higher velocity was found at 30 and 40 s after activation in sperm collected at the middle and the end of spawning period. Significantly, higher percentage of motile sperm was observed at 20, 30, and 40 s after activation in sperm sampled at the end of spawning period. This study supports the hypothesis that longer spermatozoa swim faster.

Behaviour in a standardized assay, but not metabolic or growth rate, predicts behavioural variation in an adult aquatic top predator Esox lucius in the wild.

This study tested for links among behaviour, state and life-history variables as predicted by the pace-of-life hypothesis in adult pike Esox lucius. First, a standardized open-field behavioural assay was developed to assess individual behaviour of wild-captured adult E. lucius. Behaviour within the standardized assay predicted swimming behaviour in the lake, providing an ecological validation of the assay. There was no relationship between standardized behaviour and any of the life-history and state variables, including metabolism, body condition, juvenile growth rate and adult growth rate in contrast to predictions from the pace-of-life hypothesis. This study demonstrates that it is possible to assess ecologically relevant behavioural variation in a large-bodied top predator using a standard open-field assay, but it is noteworthy that this standardized behaviour is not systematically related to standard metabolism or growth.

Fitness consequences of early life conditions and maternal size effects in a freshwater top predator.

Conditions experienced in early life stages can be an important determinant of individual life histories. In fish, environmental conditions are known to affect early survival and growth, but recent studies have also emphasized maternal effects mediated by size or age. However, the relative sensitivity of the mean fitness (population growth rate λ) to different early life impacts remains largely unexplored. Using a female-based integral projection model (IPM) parameterized from unique long-term demographic data for pike (Esox lucius), we evaluated the relative fitness consequences of different early life impacts, including (i) maternal effects of length on egg weight, potentially affecting offspring (first year) survival, and (ii) effects of temperature on offspring growth and survival. Of the seven vital rates defining the model, offspring survival could not be directly estimated and four scenarios were defined for this rate. Elasticity analyses of the IPM were performed to calculate (i) the total contribution from different lengths to the elasticity of λ to the projection kernel, and (ii) the elasticity of λ to underlying variables of female current length, female offspring length at age 1, and temperature. These elasticities were decomposed into contributions from different vital rates across length. Egg weight increased with female length, as expected, but the effect leveled off for the largest females. However, λ was largely insensitive to this effect, even when egg weight was assumed to have a strong effect on offspring survival. In contrast, λ was sensitive to early temperature conditions through growth and survival. Among mature females, the total elasticity of λ to the projection kernel generally increased with length. The results were robust to a wide range of assumptions. These results suggest that environmental conditions experienced in early life represent a more important driver of mean population growth and fitness of pike than maternal effects of size on offspring survival. We discuss two general mechanisms underlying the weak influence of this maternal effect, suggesting that these may be general for long-lived and highly fecund fishes. This model and results are relevant for the management of long-lived top predators, including many commercially important fish species.

Evolutionary divergence of adult body size and juvenile growth in sympatric subpopulations of a top predator in aquatic ecosystems.

Evolutionary theory predicts that different selective regimes may contribute to divergent evolution of body size and growth rate among populations, but most studies have focused on allopatric populations. Here, we studied five sympatric subpopulations of anadromous northern pike (Esox lucius) in the Baltic Sea subjected to allopatric habitats for a short period of their life cycle due to homing behavior. We report differences in adult body size among subpopulations that were in part due to variation in growth rate. Body size of emigrating juveniles also differed among subpopulations, and differences remained when individuals were reared in a common environment, thus indicating evolutionary divergence among subpopulations. Furthermore, a QST-FST comparison indicated that differences had evolved due to divergent selection rather than genetic drift, possibly in response to differences in selective mortality among spawning habitats during the allopatric life stage. Adult and juvenile size were negatively correlated across subpopulations, and reconstruction of growth trajectories of adult fishes suggested that body size differences developed gradually and became accentuated throughout the first years of life. These results represent rare evidence that sympatric subpopulations can evolve differences in key life-history traits despite being subjected to allopatric habitats during only a very short fraction of their life.

Photoperiodic growth enhancement in a tropical batch spawning atherinopsid, pike silverside Chirostoma estor.

The effect of photoperiod on growth and survival in early life was determined in the tropical batch spawning atherinopsid, pike silverside Chirostoma estor. The results demonstrate high sensitivity of newly hatched C. estor to photoperiod treatments up to 90 days post hatch shown by improved growth in mass (43%) under continuous illumination. This is accompanied by increased fat deposition, which suggests a critical interaction between different photoperiod-mediated mechanisms. A thorough understanding of these mechanisms can help to optimize the development of aquaculture of C. estor and similar species.

Natal departure timing from spatially varying environments is dependent of individual ontogenetic status.

Natal departure timing represents one of the first crucial decisions for juveniles born in spatially varying environments that ultimately disappear, but our knowledge on its determinants is limited. The present study aimed at understanding the determinants of juvenile natal departure by releasing individually tagged juvenile pike (Esox lucius L.) with variable body size and trophic position in a temporary flooded grassland. Specifically, we investigated whether natal departure depends on individual competitive status ('competition hypothesis'), physiological tolerance to environmental conditions ('physiological hypothesis') or individual trophic position and the spatial heterogeneity of trophic resources ('trophic hypothesis'). The results indicated that departure timing was negatively correlated with body size at release, showing that the dominance status among competing individuals was not the main trigger of juvenile departure. A positive correlation between departure timing and individual body size at departure was observed, suggesting that inter-individual variability in physiological tolerance did not explain departure patterns. While individual growth performances were similar irrespective of the timing of natal departure, stable isotope analyses revealed that juveniles with higher trophic position departed significantly earlier than individuals with lower trophic position. Therefore, the trade-off driving the use of spatially varying environments was most likely dependent upon the benefits associated with energetic returns than the costs associated with inter-individual competition or physiological stress. This result highlighted how ontogeny, and particularly ontogenetic niche shift, can play a central role in juvenile's decision to depart from natal habitats in a predatory species.

Mercury elimination by a top predator, Esox lucius.

Top-level piscivores are highly sought after for consumption in freshwater fisheries, yet these species contain the highest levels of the neurotoxin monomethylmercury (MMHg) and therefore present the greatest concern for MMHg exposure to humans. The slow elimination of MMHg is one factor that contributes to high levels of this contaminant in fish; however, little quantitative information exists on elimination rates by top predators in nature. We determined rates of MMHg elimination in northern pike (Esox lucius) by transferring fish that had naturally accumulated isotope-enriched MMHg (spike MMHg) through a whole-lake Hg loading study to a different lake. Over a period of ~7 y, pike were periodically recaptured and a small amount of muscle tissue was extracted using a nonlethal biopsy. Spike total mercury (THg) persisted in muscle tissue throughout the entire study despite discontinuing exposure upon transfer to the new lake. Spike THg burdens increased for the first ~460 d, followed by a decline to 65% of original burden levels over the next 200 d, and subsequently reached a plateau near original burden levels for the remainder of the study. We estimated the half-life of muscle THg to be 3.3 y (1193 d), roughly 1.2- to 2.7-fold slower than predicted by current elimination models. We advocate for further long-term field studies that examine kinetics of MMHg in fish to better inform predictive models estimating the recovery of MMHg-contaminated fisheries.

Inferring occurrence of growth checks in pike (Esox lucius) scales by using sequential isotopic analysis of otoliths.

RATIONALE: Sequential analysis of otolith oxygen isotope (δ(18) O(oto) ) values was performed by classical mechanical microdrilling and dual inlet mass spectrometry to infer the timing of growth checks in pike (Esox lucius) scales from Lake Annecy (France). It indicated that the first check on scales occurs before the first winter of life of the fish. METHODS: Eleven fish caught in February and June 2008 were studied. Intra-individual isotopic profiles exhibit cyclic variations that are consistent with that of modeled δ(18) O(oto) values using variations in epilimnion temperature and otolith-specific isotope fractionation equations. The positions of growth checks on scales were compared with the periodic variations of δ(18) O(oto) values on otolith radii using an allometric relationship. RESULTS: All individuals presented at least one check deposited before the first winter. Deposition of some checks might be linked to a shift in fish diet or habitat or to catch-and-release induced stress. CONCLUSIONS: Although coupling between sequential analysis of δ(18) O(oto) values and observation of scale growth features has been shown to be useful, there are limitations in the methodology, especially for the growth-attenuated region of otoliths associated with older ages.

Behavioural syndrome in a solitary predator is independent of body size and growth rate.

Models explaining behavioural syndromes often focus on state-dependency, linking behavioural variation to individual differences in other phenotypic features. Empirical studies are, however, rare. Here, we tested for a size and growth-dependent stable behavioural syndrome in the juvenile-stages of a solitary apex predator (pike, Esox lucius), shown as repeatable foraging behaviour across risk. Pike swimming activity, latency to prey attack, number of successful and unsuccessful prey attacks was measured during the presence/absence of visual contact with a competitor or predator. Foraging behaviour across risks was considered an appropriate indicator of boldness in this solitary predator where a trade-off between foraging behaviour and threat avoidance has been reported. Support was found for a behavioural syndrome, where the rank order differences in the foraging behaviour between individuals were maintained across time and risk situation. However, individual behaviour was independent of body size and growth in conditions of high food availability, showing no evidence to support the state-dependent personality hypothesis. The importance of a combination of spatial and temporal environmental variation for generating growth differences is highlighted.

Mercury and selenium concentrations in muscle tissue of different species of predatory freshwater fish and correlation between these elements.

Concentrations of total mercury and selenium were determined in 49 and 42 muscle tissue samples, respectively, of six species of predatory freshwater fish, dace (Leuciscus leuciscus), pike perch (Sander lucioperca), pike (Esox lucius), European catfish (Silurus glanis), rainbow trout (Oncorhynchus mykiss) and asp (Aspius aspius). Muscle selenium concentration did not correlate with the corresponding total mercury concentration (R² < 0.198) in all examined predatory fish species. There was an inverse correlation between the ratio Se/Hg content and the total mercury content in the muscle tissues of dace, pike perch, pike, European catfish and asp. The muscle tissue of rainbow trout exhibits a linear correlation between the ratio Se/Hg content and the total mercury content. The total mercury concentration of all examined samples did not exceed the hygienic limit for Hg for predatory fish.

Body downsizing caused by non-consumptive social stress severely depresses population growth rate.

Chronic social stress diverts energy away from growth, reproduction and immunity, and is thus a potential driver of population dynamics. However, the effects of social stress on demographic density dependence remain largely overlooked in ecological theory. Here we combine behavioural experiments, physiology and population modelling to show in a top predator (pike Esox lucius) that social stress alone may be a primary driver of demographic density dependence. Doubling pike density in experimental ponds under controlled prey availability did not significantly change prey intake by pike (i.e. did not significantly change interference or exploitative competition), but induced a neuroendocrine stress response reflecting a size-dependent dominance hierarchy, depressed pike energetic status and lowered pike body growth rate by 23 per cent. Assuming fixed size-dependent survival and fecundity functions parameterized for the Windermere (UK) pike population, stress-induced smaller body size shifts age-specific survival rates and lowers age-specific fecundity, which in Leslie matrices projects into reduced population rate of increase (lambda) by 37-56%. Our models also predict that social stress flattens elasticity profiles of lambda to age-specific survival and fecundity, thus making population persistence more dependent on old individuals. Our results suggest that accounting for non-consumptive social stress from competitors and predators is necessary to accurately understand, predict and manage food-web dynamics.

Interactions of multiple predators with different foraging modes in an aquatic food web.

Top predators can have different foraging modes that may alter their interactions and effects on food webs. Interactions between predators may be non-additive resulting from facilitation or interference, whereas their combined effects on a shared prey may result in emergent effects that are risk enhanced or risk reduced. To test the importance of multiple predators with different foraging modes, we examined the interaction between a cruising predator (largemouth bass, Micropterus salmoides) and an ambush predator (muskellunge, Esox masquinongy) foraging on a shared prey (bluegill sunfish, Lepomis macrochirus) with strong anti-predator defense behaviors. Additive and substitution designs were used to compare individual to combined predator treatments in experimental ponds. The multiple predator interaction facilitated growth of the cruising predator in the combined predator treatments, whereas predator species had substitutable effects on the growth of the ambush predator. The combined predator treatments created an emergent effect on the prey; however, the direction was dependent on the experimental design. The additive design found a risk-reducing effect, whereas the substitution design found a risk-enhancing effect for prey fish. Indirect effects from the predators weakly extended to lower trophic levels (i.e., zooplankton community). Our results highlight the need to consider differences in foraging mode of top predators, interactions between predators, and emergent effects on prey to understand food webs.

Harvest-induced disruptive selection increases variance in fitness-related traits.

The form of Darwinian selection has important ecological and management implications. Negative effects of harvesting are often ascribed to size truncation (i.e. strictly directional selection against large individuals) and resultant decrease in trait variability, which depresses capacity to buffer environmental change, hinders evolutionary rebound and ultimately impairs population recovery. However, the exact form of harvest-induced selection is generally unknown and the effects of harvest on trait variability remain unexplored. Here we use unique data from the Windermere (UK) long-term ecological experiment to show in a top predator (pike, Esox lucius) that the fishery does not induce size truncation but disruptive (diversifying) selection, and does not decrease but rather increases variability in pike somatic growth rate and size at age. This result is supported by complementary modelling approaches removing the effects of catch selectivity, selection prior to the catch and environmental variation. Therefore, fishing most likely increased genetic variability for somatic growth in pike and presumably favoured an observed rapid evolutionary rebound after fishery relaxation. Inference about the mechanisms through which harvesting negatively affects population numbers and recovery should systematically be based on a measure of the exact form of selection. From a management perspective, disruptive harvesting necessitates combining a preservation of large individuals with moderate exploitation rates, and thus provides a comprehensive tool for sustainable exploitation of natural resources.

Coexistence of behavioural types in an aquatic top predator: a response to resource limitation?

Intra-population variation in behaviour unrelated to sex, size or age exists in a variety of species. The mechanisms behind behavioural diversification have only been partly understood, but density-dependent resource availability may play a crucial role. To explore the potential coexistence of different behavioural types within a natural fish population, we conducted a radio telemetry study, measuring habitat use and swimming activity patterns of pike (Esox lucius), a sit-and-wait predatory fish. Three behavioural types co-occurred in the study lake. While two types of fish only selected vegetated littoral habitats, the third type opportunistically used all habitats and increased its pelagic occurrence in response to decreasing resource biomasses. There were no differences in size, age or lifetime growth between the three behavioural types. However, habitat-opportunistic pike were substantially more active than the other two behavioural types, which is energetically costly. The identical growth rates exhibited by all behavioural types indicate that these higher activity costs of opportunistic behaviour were compensated for by increased prey consumption in the less favourable pelagic habitat resulting in approximately equal fitness of all pike groups. We conclude that behavioural diversification in habitat use and activity reduces intraspecific competition in preferred littoral habitats. This may facilitate the emergence of an ideal free distribution of pike along resource gradients.

Assessment of larval deformities and selenium accumulation in northern pike (Esox lucius) and white sucker (Catostomus commersoni) exposed to metal mining effluent.

Uranium mining and milling operations in northern Saskatchewan (Canada) release effluents with elevated levels of certain trace metals and metalloids, including selenium. The goal of the present study was to evaluate the presence of selenium-induced deformities in northern pike (Esox lucius) and white sucker (Catostomus commersoni) larvae originating from adults collected downstream of a uranium mine. Eggs were fertilized in the field and incubated in the laboratory following a two-way (crossover) analysis-of-variance experimental design to discriminate effects from maternal transfer versus those from exposure to site water in the developing embryos. Selenium concentrations in northern pike and white sucker eggs (8.02 and 4.89 microg/g dry wt, respectively; mean +/- standard error throughout) from the exposure site were approximately two- to threefold higher than reference (2.35 +/- 0.20 and 1.94 +/- 0.25 microg/g dry wt, respectively). Among all evaluated deformities (skeletal curvatures, craniofacial deformities, fin deformities, and edema), only edema in white sucker fry from the exposure site was slightly elevated ( approximately 3%) compared to reference. The occurrence of edema, however, can be associated with factors other than selenium (e.g., other metals and organic compounds). Both fish species displayed strong linear relationships between the selenium concentrations in eggs and other tissues (muscle, liver, kidney, and bone), suggesting that selenium concentrations in eggs could be predicted from selenium concentrations in adult tissues. The lack of a clear, toxic response in the present study is in agreement with selenium thresholds for early life-stage deformities reported in other studies, with egg selenium concentrations in northern pike and white sucker collected at the exposure site being less than the 10 microg/g (dry wt) threshold associated with the presence of deformities.

Trait changes in a harvested population are driven by a dynamic tug-of-war between natural and harvest selection.

Selective harvest of large individuals should alter natural adaptive landscapes and drive evolution toward reduced somatic growth and increased reproductive investment. However, few studies have simultaneously considered the relative importance of artificial and natural selection in driving trait changes in wild populations. Using 50 years of individual-based data on Windermere pike (Esox lucius), we show that trait changes tracked the adaptive peak, which moved in the direction imposed by the dominating selective force. Individual lifetime somatic growth decreased at the start of the time series because harvest selection was strong and natural selection was too weak to override the strength of harvest selection. However, natural selection favoring fast somatic growth strengthened across the time series in parallel with the increase in pike abundance and, presumably, cannibalism. Harvest selection was overridden by natural selection when the fishing effort dwindled, triggering a rapid increase in pike somatic growth. The two selective forces appear to have acted in concert during only one short period of prey collapse that favored slow-growing pike. Moreover, increased somatic growth occurred concurrently with a reduction in reproductive investment in young and small female pike, indicating a tradeoff between growth and reproduction. The age-specific amplitude of this change paralleled the age-specific strength of harvest pressure, suggesting that reduced investment was also a response to increased life expectancy. This is the first study to demonstrate that a consideration of both natural selection and artificial selection is needed to fully explain time-varying trait dynamics in harvested populations.

Four decades of opposing natural and human-induced artificial selection acting on Windermere pike (Esox lucius).

The ability of natural selection to drive local adaptation has been appreciated ever since Darwin. Whether human impacts can impede the adaptive process has received less attention. We tested this hypothesis by quantifying natural selection and harvest selection acting on a freshwater fish (pike) over four decades. Across the time series, directional natural selection tended to favour large individuals whereas the fishery targeted large individuals. Moreover, non-linear natural selection tended to favour intermediate sized fish whereas the fishery targeted intermediate sized fish because the smallest and largest individuals were often not captured. Thus, our results unequivocally demonstrate that natural selection and fishery selection often acted in opposite directions within this natural system. Moreover, the two selective factors combined to produce reduced fitness overall and stronger stabilizing selection relative to natural selection acting alone. The long-term ramifications of such human-induced modifications to adaptive landscapes are currently unknown and certainly warrant further investigation.

Bioenergetics and growth of young-of the-year northern pike (Esox lucius) and burbot (Lota lota) exposed to metal mining effluent.

We hypothesized that exposure to metal mining effluent would reduce the ability of young-of-the-year fishes to accumulate energy reserves to survive the overwinter period (known as "winter stress syndrome") in a Canadian boreal forest watershed. Northern pike (Esox lucius) and burbot (Lota lota) were collected immediately before and after winter from a reference lake and two lakes receiving effluent. Unexpectedly, total body lipid and triglyceride, and liver triglyceride levels were greater in effluent-exposed pike and burbot in both fall and spring. However, there were no lake or season differences in growth indices of length, weight, muscle RNA/DNA ratio, or muscle protein levels in pike. In addition, total lipids and triglycerides in burbot were greater in spring compared to fall, while no seasonal differences were observed in pike, suggesting that burbot continued to feed during winter. Findings do not support the winter stress syndrome hypothesis and suggest possible direct and indirect effects of metal mining effluent on lipid dynamics of juvenile fishes.

Fluence rate or cumulative dose? Vulnerability of larval northern pike (Esox lucius) to ultraviolet radiation.

Newly hatched larvae of northern pike were exposed in the laboratory to four fluence rates of ultraviolet radiation (UVR; 290-400 nm) over three different time periods, resulting in total doses ranging from 3.0 +/- 0.2 to 63.0 +/- 4.4 kJ.m(-2). Mortality and behavior of the larvae were followed for 8-12 days, and growth measured at the end of the experiment. Also, the principle of reciprocity-that the UVR-induced mortality depends on the cumulative dose, independent of fluence rate-was tested. Fluence rates higher than 1480 +/- 150 mW.m(-2) caused mortality and growth retardation. The highest fluence rate (3040 +/- 210 mW.m(-2)) caused 100% mortality in 5 days. All fluence rates caused behavioral disorders, which led to death at fluence rates higher than 1480 mW.m(-2). Reciprocity failure occurred with the lowest and highest dose (550 +/- 45 and 3040 +/- 210 mW.m(-2), respectively). The results show that fluence rate is of primary importance when assessing the UVR-related risk.