Adaptive divergence in body size overrides the effects of plasticity across natural habitats in the brown trout.

The evolution of life-history traits is characterized by trade-offs between different selection pressures, as well as plasticity across environmental conditions. Yet, studies on local adaptation are often performed under artificial conditions, leaving two issues unexplored: (i) how consistent are laboratory inferred local adaptations under natural conditions and (ii) how much phenotypic variation is attributed to phenotypic plasticity and to adaptive evolution, respectively, across environmental conditions? We reared fish from six locally adapted (domesticated and wild) populations of anadromous brown trout (Salmo trutta) in one semi-natural and three natural streams and recorded a key life-history trait (body size at the end of first growth season). We found that population-specific reaction norms were close to parallel across different streams and Q ST was similar - and larger than F ST - within all streams, indicating a consistency of local adaptation in body size across natural environments. The amount of variation explained by population origin exceeded the variation across stream environments, indicating that genetic effects derived from adaptive processes have a stronger effect on phenotypic variation than plasticity induced by environmental conditions. These results suggest that plasticity does not "swamp" the phenotypic variation, and that selection may thus be efficient in generating genetic change.

Strong divergence in trait means but not in plasticity across hatchery and wild populations of sea-run brown trout Salmo trutta.

There is ample evidence that organisms adapt to their native environment when gene flow is restricted. However, evolution of plastic responses across discrete environments is less well examined. We studied divergence in means and plasticity across wild and hatchery populations of sea-run brown trout (Salmo trutta) in a common garden experiment with two rearing environments (hatchery and a nearly natural experimental stream). Since natural and hatchery environments differ, this arrangement provides an experiment in contemporary adaptation across the two environments. A Q(ST) - F(ST) approach was used to investigate local adaptation in survival and growth over the first summer. We found evidence for divergent selection in survival in 1 year and in body length in both years and rearing environments. In general, the hatchery populations had higher survival and larger body size in both environments. Q(ST) in body size did not differ between the rearing environments, and constitutive divergence in the means was in all cases stronger than divergence in the plastic responses. These results suggest that in this system, constitutive changes in mean trait values are more important for local adaptation than increased plasticity. In addition, ex situ rearing conditions induce changes in trait means that are adaptive in the hatchery, but potentially harmful in the wild, suggesting that hatchery rearing is likely to be a suboptimal management strategy for trout populations facing selection in the stream environment.

Bioenergy potential of Ulva lactuca: biomass yield, methane production and combustion.

The biomass production potential at temperate latitudes (56°N), and the quality of the biomass for energy production (anaerobic digestion to methane and direct combustion) were investigated for the green macroalgae, Ulva lactuca. The algae were cultivated in a land based facility demonstrating a production potential of 45T (TS) ha(-1) y(-1). Biogas production from fresh and macerated U. lactuca yielded up to 271 ml CH(4) g(-1) VS, which is in the range of the methane production from cattle manure and land based energy crops, such as grass-clover. Drying of the biomass resulted in a 5-9-fold increase in weight specific methane production compared to wet biomass. Ash and alkali contents are the main challenges in the use of U. lactuca for direct combustion. Application of a bio-refinery concept could increase the economical value of the U. lactuca biomass as well as improve its suitability for production of bioenergy.

Frequency distribution of coping strategies in four populations of brown trout (Salmo trutta).

In a challenging situation some animals respond by active avoidance, aggression and an activation of the sympathetic nervous system whereas others respond by immobility, low levels of aggression and a predominant adrenocortical stress response. When consistent over time and across situations such inter-individual differences in behavioural and physiological stress responses are referred to as stress coping strategies. In a previous study we reported the existence of two distinct stress coping strategies in a sea-ranched brown trout (Salmo trutta) population. Using the same method, we here show that four brown trout populations with different origin, but reared under identical conditions, differ in their endocrine stress response, behaviour during hypoxia and aggression. Further more, if individuals are classified as high- and low responsive based on post-stress blood plasma noradrenalin levels (indicator of sympathetic reactivity) the frequency distribution shows that populations with hatchery origin are biased towards having higher frequencies of high responsive individuals. However, the number of high responsive trout ranges from 14-48% in the different populations which shows that generally the frequency is biased towards lower levels of high responsive individuals. We discuss different frequency-dependent mechanisms that maintain multiple phenotypes in populations and speculate about differences in selection regime among the studied populations.

Developmental responses to predation risk in morphologically defended mayflies.

Densities and species composition of predators could affect morphological defences, larval development and the timing of emergence of their prey. To address this issue we studied the morphology and life history of an ephemerellid mayfly, Ephemerella invaria, from two streams in a deciduous forested drainage basin in central New York. Both streams contained predatory fish, but densities and species composition of fish differed. A field survey provided evidence that Ephemerella inhabiting a stream with > or =10 fish species and high relative densities of fish emerged several weeks earlier and at smaller sizes than Ephemerella inhabiting a nearby tributary with approximately 2 fish species and low relative densities of fish. However, the two populations of mayflies showed no differences in defensive morphology or growth rates. In laboratory rearing experiments, we exposed Ephemerella larvae from these two locations to fish chemical cues or control water (no fish) over 2 months to test whether differences in life histories could be attributed to fish. Fish cues induced faster larval development, but also smaller size of mature Ephemerella individuals from both high and low predator locations. Although shorter development times in more dangerous environments could increase larval survival, smaller size of females results in a fecundity cost associated with this life history shift. Consistent with the field studies, laboratory rearing experiments revealed no effects of fish cues on Ephemerella's morphological defences. These data suggest that variation in the density or species composition of predators may favour the evolution of developmental plasticity to reduce mortality in the larval environment.

Effects of a benthivorous and a drift-feeding fish on a benthic stream assemblage.

I assessed the impact of both drift-feeding (Salmo trutta, brown trout) and benthic-feeding (Cottus gobio, bullhead) fish on a benthic assemblage during a 1-month-long field experiment. I used enclosures containing cobble/gravel substrata with 6-mm mesh net that allowed invertebrates to drift freely in and out of enclosures. Four treatments, arranged in a factorial design, were tested: a predator-free control, bullheads only (2.67 bullheads/m2, two per enclosure), brown trout only (2.67 brown trout/m2, two per enclosure), and bullheads and brown trout together (2.67 fish/m2, one of each). Bullheads reduced the densities of seven invertebrate taxa (Gammarus pulex amphipods, Baetis rhodani mayfly nymphs, Leuctra spp. stonefly nymphs, Polycentropus spp. caddis larvae, Pacifastacus leniusculus signal crayfishes, Simuliidae, blackfly larvae, and Limnephilidae, caddis larvae) whereas brown trout only affected one taxon (B. rhodani). The weaker effect of brown trout on benthic prey was probably related to its heavy reliance on terrestrial prey. Dietary analyses showed that more than 80% of prey consumed by brown trout were terrestrial animals, whereas bullhead only consumed benthic prey. Neither bullhead nor brown trout affected the absolute number of immigrating or emigrating invertebrates in enclosures, but bullhead affected the per capita emigration rates of G. pulex. Direct predation by bullhead was more important than avoidance behaviour (drift) in determining densities of six of the seven taxa; only G. pulex densities were equally affected by avoidance behaviour and direct predation. Direct predation by brown trout was also more important in determining densities of B. rhodani. The presence of bullhead raised periphyton biomass, presumably mediated via their consumption of grazers. Brown trout had no effect on periphyton biomass and these results indicate that the presence of alternative prey, in this case terrestrial animals, may have repercussions for fish-benthic macroinvertebrate-periphyton interactions and may potentially explain some of the variable impacts of fish on benthic macroinvertebrates that have been reported in the literature.

The impact of vertebrate and invertebrate predators on a stream benthic community.

I assessed the impact of both vertebrate and invertebrate predators on a lotic benthic community in a 1-month-long experiment, using enclosures containing cobble/gravel bottoms, with large-mesh netting that allowed invertebrates to drift freely. Brown trout (Salmo trutta) and leeches (Erpobdella octoculata) were used as predators and four treatments were tested: a predator-free control, leeches only, trout only, and leeches and trout together. A density of 26.7 leeches/m2 (20 leeches/enclosure) and 1.3 trout/m2 (one trout per enclosure) was stocked into the enclosures. The total biomass of invertebrate prey was significantly lower in the trout and trout plus leech treatments than in the leech and control treatments, which were due to strong negative effects of trout on Gammarus. On the individual prey taxon level, both trout and leeches affected the abundance of Asellus , Baetis and Ephemerella, whereas the abundance of Gammarus was only affected by trout, and the abundance of Orthocladiinae and Limnephilidae was only affected by leeches. In the treatment with trout and leeches together, the abundance of Ephemerella and Baetis was higher than when trout or leeches were alone, which was probably due to predator interactions. Leeches and trout had no effects on prey immigration but did affect per capita emigration rates. Both trout and leeches indirectly increased periphyton biomass in enclosures, probably due to their strong effects on grazers. Both trout and leeches were size-selective predators, with trout selecting large prey, and leeches selecting small prey. Size-selective predation by trout and leeches affected the size structure of five commonly consumed prey taxa. Trout produced prey populations of small sizes owing to consumption of large prey as well as increased emigration out of enclosures by these large prey. Leech predation produced prey assemblages of larger size owing to consumption and increased emigration of small prey. These results suggest that in lotic habits, predatory invertebrates can be as strong interactors as vertebrate predators.