The cost of catching up: increased winter mortality following structural growth compensation in the wild

Although laboratory and observational studies suggest that many animals are capable of compensatory growth after periods of food shortage, few field experiments have demonstrated structural growth compensation in the wild. Here, we addressed the hypotheses that (i) food restriction can induce structural compensatory growth in free-living animals, (ii) that compensation is proportional to the level of body size retardation and (iii) that compensation induces mortality costs. To test these, wild brown trout (Salmo trutta) yearlings were brought to the lab, tagged individually, subjected to four levels of food deprivation (including a control), released back into the native stream and recaptured after one, five and ten months. Brown trout fully restored condition and partially restored mass within a month, whereas compensation in structure (i.e. body length) was not evident until after five months, supporting hypothesis 1. As the level of growth compensation was similar among the three deprived groups, hypothesis 2 was not supported. A final recapture after winter revealed delayed mortality, apparently induced by the compensatory response in the deprived groups, which is consistent with hypothesis 3. To our knowledge, this is the first field experiment demonstrating structural compensatory growth and associated costs in a wild animal population.     

Compensatory growth offsets poor condition in native trout populations

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Compensatory growth for free? A field experiment on brown trout,Salmo trutta

Laboratory studies suggest that animals may be capable of compensatory growth after periods of food shortage. There is, however, a lack of field experiments investigating the incidence and consequences of compensatory growth in the wild, and the relevance of compensatory responses in natural populations has recently been questioned. Here we addressed the hypotheses that (1) food restriction during critical growth periods can induce compensatory growth, and (2) that compensatory growth is associated with delayed costs in natural populations. These hypotheses were addressed by (1) manipulating the food intake of brown trout in spring, (2) measuring growth rate responses over the first month following release, and (3) measuring growth and mortality (i.e. recapture rate) over the subsequent fall and winter. We found that brown trout restored lost body weight and condition within a month, providing the first experimental demonstration of compensatory growth in the wild. However, no delayed costs of the compensatory response could be detected within the timespan of the experiment. We suggest that wild brown trout have an adapted "buffer capacity" to withstand fluctuations in food supply, allowing restoration of lost lipid reserves when feeding conditions improve. However, when prolonged food deprivation affect structural components, compensation may not be possible without compromising long-term performance.     

Stocking hatchery‐reared brown trout in different densities into a wild population – a comparison of growth and movement

In spring 2001 and 2002 a small stream was stocked with tagged hatchery‐reared yearling brown trout ( Salmo trutta ), in order to study their influence on the resident brown trout population. The stream was separated into six sections: two sections without stocking, two sections where stocking doubled the trout population and two sections where the fish population was quadrupled. The working hypothesis was that due to food limitation (competition) growth of the wild fish will be negatively influenced by stocking, and wild fish will be displaced by the (possibly more aggressive) hatchery fish. Surprisingly, growth rate of wild and stocked fish of the same age was similar and independent of stocking density. Two main reasons may be responsible for this finding: only a low percentage of the stocked fish remained in the stream, and food was not limited during summer. Only 12–19% of the stocked fish were recaptured after six months, in contrats to 40–70% of one‐year old and up to 100% of older wild trout. The wild fish were not displaced by hatchery‐reared fish: During summer the wild fish remained more or less stationary, whereas most of the stocked trout had left their release site. The results indicate that in a natural stream stocking of hatchery reared brown trout does not influence negatively growth and movement of the wild fish independent of stocking density.     

Density‐dependent growth in hatchery‐reared brown trout released into a natural stream

Hatchery‐reared brown trout Salmo trutta stocked in a natural stream in addition to resident wild brown trout grew more slowly than those stocked with an experimentally reduced density of brown wild trout. In both cases, hatchery‐reared brown trout grew more slowly than resident wild fish in control sections. Mortality and movements did not differ among the three categories of fish. The results showed that growth of stocked hatchery‐reared brown trout parr was density‐dependent, most likely as a consequence of increased competition. Thus, supplementary release of hatchery‐reared fish did not necessarily increase biomass.     

Environmentally induced spatio-temporal variations in the fecundity of brown trout Salmo trutta L.: trade-offs between egg size and number

1. Resident brown trout Salmo trutta in the Esva River basin (north Spain) live in a patchy environment with tracts of riparian forest or meadow along stream banks. This study assessed whether the reproductive traits of brown trout from four contrasting sites reflected site-specific factors.
2. Length at maturity (10.5–11 cm of 1 + individuals) was the same in the four sites examined but slowest growers in slow-growing sub-populations delayed maturity for 1 year relative to fast-growing fish. The analysis of monthly variations in egg size and number suggest that two 'decisions' in two consecutive years are required to complete spawning. The first concerns the number of eggs, determined when trout are still 0 +, and the second concerns egg size.
3. At three sites, egg size and number did not differ significantly between years but highly significant interannual variations were apparent at another site. Fish length was the major determinant of egg size and number at all sites but for any given length, brown trout at sites where the fish exhibited higher growth rates spawned more, but smaller, eggs than those at slow-growing sites. This spatial pattern was identical to the temporal pattern exhibited by trout at another site. The combination of temporal (year-to-year) and spatial (between rivers) variations in egg size and number showed a significant negative correlation, supporting the operation of a trade-off between these two traits.
4. The trade-off between egg size and number seems to be determined by site-specific factors, with slow-growing trout at sites which are fully covered by canopy spawning fewer, but larger, eggs than fast-growers in unshaded sites.     

BRIEF COMMUNICATIONS Persistence times of four amphipod species in the stomachs of brown trout

Four amphipod species fed to brown trout Salmo trutta persisted in an easily identifiable form in trout stomachs for similar lengths of time. The data also allowed estimation of the maximum length of time ingested amphipods could persist in an identifiable form in brown trout stomachs at different temperatures. The data are thus useful in assessment of fish feeding patterns on amphipod assemblages in the laboratory and field.     

Brown trout and food web interactions in a Minnesota stream

1. We examined indirect, community‐level interactions in a stream that contained non‐native brown trout (Salmo trutta Linnaeus), native brook trout (Salvelinus fontinalis Mitchill) and native slimy sculpin (Cottus cognatus Richardson). Our objectives were to examine benthic invertebrate composition and prey selection of fishes (measured by total invertebrate dry mass, dry mass of individual invertebrate taxa and relative proportion of invertebrate taxa in the benthos and diet) among treatments (no fish, juvenile brook trout alone, juvenile brown trout alone, sculpin with brook trout and sculpin with brown trout). 2. We assigned treatments to 1 m² enclosures/exclosures placed in riffles in Valley Creek, Minnesota, and conducted six experimental trials. We used three designs of fish densities (addition of trout to a constant number of sculpin with unequal numbers of trout and sculpin; addition of trout to a constant number of sculpin with equal numbers of trout and sculpin; and replacement of half the sculpin with an equal number of trout) to investigate the relative strength of interspecific versus intraspecific interactions. 3. Presence of fish (all three species, alone or in combined‐species treatments) was not associated with changes in total dry mass of benthic invertebrates or shifts in relative abundance of benthic invertebrate taxa, regardless of fish density design. 4. Brook trout and sculpin diets did not change when each species was alone compared with treatments of both species together. Likewise, we did not find evidence for shifts in brown trout or sculpin diets when each species was alone or together. 5. We suggest that native brook trout and non‐native brown trout fill similar niches in Valley Creek. We did not find evidence that either species had an effect on stream communities, potentially due to high invertebrate productivity in Valley Creek.     

Growth, carcass composition and plasma growth hormone levels in cyclically fed rainbow trout

Growth, body composition and plasma growth hormone levels were recorded weekly for 24 weeks in rainbow trout Oncorhynchus mykiss . Underyearling rainbow trout were individually identified using coded tags and placed on either a cyclic feeding regime of 3 weeks of deprivation followed by 3 weeks of feeding or a daily feeding regime. No significant difference was found in standard length and mass among the cyclically fed and daily fed fish at the end of the experiment. For cyclically fed fish, the absolute specific growth rate and condition factor reached a maximum during the last week of refeeding. Cyclically fed fish had a significantly higher moisture and protein content and lower lipid levels relative to fish fed daily. Absolute mass and fat loss in the deprivation phase of the feeding cycle decreased in intensity with subsequent feeding cycles, indicating that the fish were acclimatizing to the feeding regime. It was proposed that this response was an adaptation against possible adverse effects in the adults ( e.g. locomotor performance, bone ossification rates, fat deposition rate, growth rate and age at sexual maturity). Plasma growth hormone concentrations were not affected by cyclic feeding indicating that variations in plasma growth hormone concentration are not the cause of compensatory growth in rainbow trout.     

Synchrony in brown trout, Salmo trutta, population dynamics: a 'Moran effect' on early-life stages

Synchrony among populations (i.e. spatial covariation in temporal fluctuations of population size or growth rate) is a common feature to many animals. Both large-scale autocorrelated climatic factors (the 'Moran effect') and dispersal between populations are candidates to explain synchrony, although their relative influence is difficult to assess. Only a few investigations have reported patterns of synchrony among freshwater populations, and even fewer directly related these patterns to an environmental variable. In the present study, we analysed the spatio-temporal patterns of fluctuation of 57 brown trout populations widespread across France, each sampled continuously during 5 years. We compared the respective influence of connectivity and stream distance within basins (i.e. that potentially allow a basin-scale dispersal) and environmental factors (hydrological and air temperature variables, available for 37 sites) on the synchrony of brown trout cohort densities (0+, 1+ and adults). A series of Mantel tests revealed that the degree of synchrony was not related to connectivity or stream distance between sites, suggesting no effect of dispersal at the basin-scale. The degree of synchrony among sites for the 0+ fish was significantly related to the degree of hydrological synchrony (based on high flows during the emergence period). For all three age classes, the synchrony in the temperature patterns did not explain synchrony in trout dynamics. Our results allow us to discuss the respective influence of dispersal and climatic factors on the spatio-temporal patterns of trout dynamics at the basin scale.     

Juvenile migration in brown trout: a consequence of energetic state

1. We explored the mechanisms determining age and size at juvenile migration in brown trout Salmo trutta L. A ¹³³Cs tracer methodology was used to estimate food consumption of juvenile brown trout in a Norwegian stream, and the energy budgets of early migrants and stream residents were compared.
2. Fast-growing brown trout migrated to the lake earlier and at a smaller body size than slower-growing individuals. The 2+ migrants were significantly larger than those that remained 1 or more years longer in the stream. The 3+ migrants were significantly larger than the 2+ migrants. Some fast-growing males matured in the stream, whereas all females left the stream before maturing sexually.
3. The food consumption and the energy budgets for 2+ migrants were more than four times higher than those of the resident 2+ fish. Total energy allocated to growth was also higher among migrants, and the total metabolic costs were five times higher among migrants than among resident fish.
4. The proportional energy allocation to growth among the 2+ migrants was much lower (about half) than that of those remaining longer in the stream. The reduction in the proportion of energy available for growth from age 1+ to 2+ was larger among migrants (88%) than among resident fish (68%). Reduction in the proportion of energy available for growth is a probable explanation for why migrations are initiated at age 2.
5. Our study supports the hypothesis that fast-growing individuals shift their niche earlier and at a smaller body size than slower-growing individuals because they maintain higher metabolic rates and are energetically constrained at a younger age by limited food resources than slow growers.     

Prior residence, aggression and territory acquisition in hatchery-reared and wild brown trout

In an artificial stream environment, established wild brown trout initiated 44% of the mean aggressive acts whilst hatchery-reared trout initiated 34% and introduced wild trout initiated 22%. Established wild fish maintained home stations closer to a point source of feed than did both hatchery-reared and introduced wild conspecifics. Established wild fish were the only group to show a positive mean specific growth rate during the trials. Introduced wild fish showed a slightly negative mean specific growth rate, whilst introduced hatchery-reared fish exhibited a considerable negative mean specific growth rate. These results suggest that established wild brown trout in a semi-natural stream environment display a prior-resident effect over late introductions of hatchery-reared and wild conspecifics. Introduced hatcheryreared fish were more aggressive and exhibited a lower mean specific growth rate than simultaneously stocked wild fish, suggesting that excessive expenditure of energy for unnecessary aggression may contribute to the poor survival of hatchery-reared fish after they are stocked into streams.     

Long-term population demographics of native brook trout following manipulative reduction of an invader

Although laboratory studies have provided evidence for negative interactions between brook trout and brown trout, it is unknown how these interactions affect larger scale demographics in a natural setting. We tested the effects of invasive brown trout on brook trout demographics by removing brown trout from a sympatric population using a before–after control-impact study design. The study was conducted across a large stream network for a period of 6 years. Abundance of brook trout increased after brown trout removal primarily as a result of increased recruitment and immigration. Size structure also shifted towards larger individuals as a result of increased growth rates and a decrease in emigration of larger trout. Size at maturity and body condition did not change after brown trout removal. Adult brook trout survival increased during the post-treatment period in both the treatment and control reach. A decrease in flood intensity during the post-treatment time period may have led to increased survival. Adult survival may not be the best metric to use when assessing interactions between trout species, especially when the subordinate species has suitable areas to emigrate.     

Ultrastructural biomarkers as tools to characterize the health status of fish in contaminated streams

Ultrastructural biomarkers in gill, liver, andkidney of brown trout (Salmo truttaf. fario) and stone loach (Barbatulabarbatula) were investigated over afive year period. The cellular damage of theorgans was assessed semi-quantitatively basedon a three-step classification ofultrastructural responses. Data obtained forfish exposed under semi-field conditions in twodifferently polluted test streams and in thelaboratory demonstrated that the ultrastructureof the organs can be correlated with differentpollutant exposure conditions. Cellular damagewas generally more severe in fish exposed to acomplexly polluted stream than in those exposedto a moderately polluted stream or to tap waterin the laboratory. Histopathological effects inliver and gill of trout were more pronounced inferal fish than in transplanted fish, whereasresponses in the kidney in both species, and inliver and gill of loach, were similar forintroduced and feral fish. In a laboratoryexperiment where trout were exposed todifferent mixtures of pollutants includingpesticides, PAH, and ammonia, only theultrastructure of kidney and liver showedsignificant differences between the threeexperimental set-ups. In a recovery experiment,where trout were transferred from thesemi-field condition back to the laboratory,ultrastructural investigations showed adifferential capacity of the respective organsto recover from stress under field conditions.Kidney and liver fully recovered after threemonths under control condition whereas gillsdemonstrated only partial recovery.     

Non-equivalence of growth arrest induced by predation risk or food limitation: context-dependent compensatory growth in anuran tadpoles

1. To gain insight into the evolution of compensatory growth, we studied the growth patterns of anuran (Rana temporaria) larvae following either a period of exogenous growth depression (food restriction) or a period of endogenous depression (exposure to predators). We also investigated the potential deferred costs that larval compensatory growth could impose on post-metamorphic individuals. 2. Food-deprived larvae exhibited full compensatory growth in response to reduced growth rates caused by food limitation, and the growth trajectories of low- and high-rations tadpoles converged before the onset of metamorphosis. 3. According to our predictions, individuals exposed to larval predators did not show growth compensation following predator removal despite undergoing a significant reduction in growth rate associated with low activity levels. 4. Jumping ability of individuals exposed to predators during only 20 days from the commencement of the larval phase was equivalent to that of non-exposed animals, and greater than the jumping capacity of those maintained with predators until the time of metamorphosis. This pattern was consistent with the pattern observed for variation in relative leg length. 5. These results support the suggestion that submaximum and compensatory growth could have evolved to minimize the overall growth/mortality costs in environments with high spatiotemporal variation in predation intensity.     

Growth‐rate variation in brown trout in small neighbouring streams: evidence for density‐dependence?

The within- and among-population variation in individual growth rate of brown trout Salmo trutta L. was studied in five small neighbouring streams (seven isolated populations) within a distance of 70 km in east Norway. Observed growth rate was only weakly correlated with predicted maximum growth rate based on laboratory models, and there was a significant interaction with site. A generalized linear model showed that growth rate was positively correlated with temperature, but also that growth rate decreased as the summer season progressed. This might indicate either a seasonal decline in food availability or appetite, or a change in energy allocation strategy. Growth rate decreased with increasing fish age, probably as an effect of sexual maturation of older fish, and differential allocation of protein and lipid among different size-groups of brown trout. After adjusting for variation in temperature, season, and fish mass, there was still significant among-site variation in growth rate. A significant part of this variation was due to variation in brown trout density and the presence or absence of Alpine bullhead Cottus poecilopus . Growth rate decreased with increasing brown trout density, and was lower in the presence than in the absence of Alpine bullhead after correcting for variation in brown trout density. This last result may indicate the presence of interspecific competition.     

The fish populations of some streams in Wales and northern England in relation to acidity and associated factors

Electrofishing and water quality surveys were carried out on 60 upland moorland streams in central and north Wales and the Peak District of England (south Pennines). A number of biological characteristics of the fish populations were recorded, including species representation, population density and biomass and, for brown trout, Salmo trutta L., only, growth, condition factor and diet. The stream waters were also analysed over a 2-year period for pH, calcium, aluminium and the heavy metals copper, zinc and lead.
Mean pH levels in the streams ranged from 5.0 to 7.6, with calcium concentrations in the range 0.80–27 mg 1⁻¹. Trace metal concentrations (Al, Cu, Zn, Pb) tended to be elevated in the more acid streams. The latter had fewer fish species, and a higher proportion was fishless, 73% at pH < 5.5 compared with 6% at pH >6.5. Trout and eels, Anguilla anguilla L., predominated at lower pH values, but at reduced population levels.
Dietary composition and stomach fullness of trout were similar throughout the stream pH range. More than 40% of the diet was derived from surface and terrestrial sources, supplementing autochthonous items. Growth (as back-calculated length to end of first year of life) and condition factor did not vary significantly with respect to stream acidity. It is thus concluded that the impoverishment of trout populations in the more acid streams was not a second-order effect acting through the lower trophic levels.
Water quality acting directly on fish would appear to account for the poor status of salmonid fisheries in acid streams. Absence or paucity of salmonids was related to high levels of labile monomeric aluminium (>40 μgl⁻¹) or Cu-Zn-Pb toxicity (>0.4 t-LC50 to rainbow trout, Salmo gairdneri Richardson). Limiting values in this study correspond well with published laboratory and field data.     

The influence of large wood on brown trout (Salmo trutta) behaviour and surface foraging

1. Changes in riparian vegetation owing to forest harvesting may affect the input of large wood, a major structural element, to streams. Studies of large wood impacts on stream fish have focused on population‐level responses, whereas little attention has been given to how wood affects fish behaviour. 2. In a laboratory stream experiment, we tested how two size classes of brown trout, Salmo trutta, (mean size of 85 and 125 mm), alone and together, responded to a gradient of large wood in terms of activity, foraging on terrestrial drift and interactions between conspecifics. 3. The results showed that the presence of large wood significantly reduced the overall activity of the fish, the number of agonistic interactions between individuals and the proportion of captured prey. However, activity decreased relatively more than the proportion of captured prey, resulting in a significant positive net effect of wood on the number of prey captures per time spent active (PTA). This indicates that trout living in habitats with high wood density may have a higher net energy gain than trout living in habitats with less wood. 4. There were no observable size‐class differences in the benefits of large wood or in the utilisation of surface‐drifting terrestrial prey. 5. These results suggest that the presence of large wood may be an important factor shaping stream communities and that a lack of structural complexity may decrease energy gain, increase agonistic interactions and, consequently, lower the production of brown trout.     

Size-independent growth in fishes: patterns, models and metrics

A combination of a dynamic energy budget (DEB) model, field data on Atlantic salmon Salmo salar and brown trout Salmo trutta and laboratory data on Atlantic salmon was used to assess the underlying assumptions of three different metrics of growth including specific growth rate (G), standardized mass‐specific growth rate (G_S) and absolute growth rate in length (G_L) in salmonids. Close agreement was found between predictions of the DEB model and the assumptions of linear growth in length and parabolic growth in mass. Field data comparing spring growth rates of age 1+ year and 2+ year Atlantic salmon demonstrated that in all years the larger age 2+ year fish exhibited a significantly lower G, but differences in growth in terms of G_S and G_L depended on the year examined. For brown trout, larger age 2+ year fish also consistently exhibited slower growth rates in terms of G but grew at similar rates as age 1+ year fish in terms of G_S and G_L. Laboratory results revealed that during the age 0+ year (autumn) the divergence in growth between future Atlantic salmon smolts and non‐smolts was similar in terms of all three metrics with smolts displaying higher growth than non‐smolts, however, both G_S and G_L indicated that smolts maintain relatively fast growth into the late autumn where G suggested that both smolts and non‐smolts exhibit a sharp decrease in growth from October to November. During the spring, patterns of growth in length were significantly decoupled from patterns in growth in mass. Smolts maintained relatively fast growth though April in length but not in mass. These results suggest G_S can be a useful alternative to G as a size‐independent measure of growth rate in immature salmonids. In addition, during certain growth stanzas, G_S may be highly correlated with G_L. The decoupling of growth in mass from growth in length over ontogeny, however, may necessitate a combination of metrics to adequately describe variation in growth depending on ontogenetic stage particularly if life histories differ.     

Characteristics of the spawning migrations of brown trout, Salmo trutta L., and rainbow trout, S. gairdneri Richardson, in Great Lake, Tasmania

Upstream spawning migrations of mature brown trout, S. trutta , and rainbow trout, S. gairdneri , were studied in Liawenee Canal, Great Lake from 1949 to 1985. Brown trout migrations normally occurred from early April to mid-May and rainbow trout from late August to early November. In 1983, 16 425 brown trout and 1338 rainbow trout passed through a fixed upstream diversion trap. Brown trout spawning migrations occurred predominantly over the temperature range 6–10° C, while rainbow trout migrated predominantly over the range 5–11° C. Migrations peaked at water temperatures of 7.6°C (males) and 7.8°C (females) for brown trout, and 8.3°C (males) and 9.6°C (females) for rainbow trout. Rainbow trout migrations occurred at high flow conditions and were positively correlated with canal flow increases, while brown trout migrated under low canal flow. Mean length, weight and condition of rainbow trout of both sexes decreased significantly during migrations. Female brown trout decreased in weight and condition but not in length; male brown trout did not change in condition despite decreases in both length and weight during migrations. Overall sex ratio was 2:1 (female:male) for both species, with the relative proportion of male fish decreasing as migrations progressed. Age composition changed during migrations; dominant age classes were 3 < 4 < 5 + years for both species. Comparison of length, weight, condition and age revealed minor changes during the 37-year period 1949–1985.