Karl Filipsson, Eva Bergman, Larry Greenberg, Martin Österling, Johan Watz and Ann Erlandsson recently published the paper “Temperature and predator-mediated regulation of plasma cortisol and brain gene expression in juvenile brown trout (Salmo trutta)” in the journal Frontiers in Zoology.

In this study, we tested how temperature and the presence of a cold-water adapted predatory fish (burbot, Lota lota) affected primary stress responses (i.e. cortisol and mRNA levels of stress-related genes) in juvenile brown trout (Salmo trutta). We found that trout had elevated cortisol levels in the presence of burbot, and that stress-related gene expressions varied a lot with temperature. In addition, we found that the predator-induced effects on mRNA levels were temperature dependent for some of the genes. This, together with the directly temperature-mediated effects that we observed in our study, suggest that warming winters can have major impact on primary stress responses in overwintering salmonids, for instance in encounters with predators.

In the abstract of the paper, we wrote that:

“Temperature affects many aspects of performance in poikilotherms, including how prey respond when encountering predators. Studies of anti-predator responses in fish mainly have focused on behaviour, whereas physiological responses regulated through the hypothalamic-pituitary-interrenal axis have received little attention. We examined plasma cortisol and mRNA levels of stress-related genes in juvenile brown trout (Salmo trutta) at 3 and 8 °C in the presence and absence of a piscivorous fish (burbot, Lota lota).

One of the experimental aquaria used for the study.

A redundancy analysis revealed that both water temperature and the presence of the predator explained a significant amount of the observed variation in cortisol and mRNA levels (11.4 and 2.8%, respectively). Trout had higher cortisol levels in the presence than in the absence of the predator. Analyses of individual gene expressions revealed that trout had significantly higher mRNA levels for 11 of the 16 examined genes at 3 than at 8 °C, and for one gene (retinol-binding protein 1), mRNA levels were higher in the presence than in the absence of the predator. Moreover, we found interaction effects between temperature and predator presence for two genes that code for serotonin and glucocorticoid receptors.

We extracted mRNA from the forebrain (telencephalon) of the trout. The picture shows a trout brain after dissection, where the telencephalon is the two upper lobes.

Our results suggest that piscivorous fish elicit primary stress responses in juvenile salmonids and that some of these responses may be temperature dependent. In addition, this study emphasizes the strong temperature dependence of primary stress responses in poikilotherms, with possible implications for a warming climate.”

You can read the paper for free on the journal website, as the paper is published open access through funding provided by Karlstad University.

Lutz Eckstein, professor at Karlstad University, together with Yves Klinger, David HorlemannAnnette Otte and Kristin Ludewig recently published the paper Germination of the invasive legume Lupinus polyphyllus depends on cutting date and seed morphology in the journal NeoBiota. The paper is accessible open access on the journal website, or you can read a summary of the paper below:

In semi-natural grasslands, mowing leads to the dispersal of species that have viable seeds at the right time. For invasive plant species in grasslands, dispersal by mowing should be avoided, and information on the effect of cutting date on the germination of invasive species is needed. We investigated the germination of seeds of the invasive legume Lupinus polyphyllus Lindl. depending on the cutting date. We measured seed traits associated with successful germination that can be assessed by managers for an improved timing of control measures. Germination patterns were highly asynchronous and differed between seeds cut at different dates. Seeds cut early, being green and soft, tended to germinate in autumn. Seeds cut late, being dark and hard, were more prone to germinate the following spring, after winter stratification. This allows the species to utilize germination niches throughout the year, indicating a bet-hedging strategy.

Lupine seeds used in the study

Seed color and the percentage of hard seeds were good predictors of germination percentage. Managers should prevent the species from producing black and hard seeds, while cutting plants carrying green and soft seeds appears less problematic. Furthermore, germination patterns differed between experiments in climate chambers and in the common garden, mainly because germination of dormant seeds was lower in climate chambers. We propose that more germination experiments under ambient weather conditions should be done, as they can give valuable information on the germination dynamics of invasive species.

Johan Watz, David Aldvén, Antonis Apostolos Brouziotis, Niclas Carlsson, Eirini Karathanou, Kristine Lund‐Bjørnås, Gustav Lundqvist, Martin Österling, John J. Piccolo and Olle Calles recently published the paper “Social behaviour of European grayling before and after flow peaks in restored and unrestored habitats” in the journal River Research and Applications.

In the abstract of the paper the authors write:

“Cost‐effective implementation of fish‐friendly hydropower flow operation and habitat restoration measures require an understanding of their effects on fitness‐related behaviours of stream fish. Here, we investigated how changes in flow and bottom structure influence the social behaviour of European grayling, using large experimental flumes (700 L s−1), with and without added boulders (i.e., restored and unrestored habitat). Grayling increased their distance to nearest neighbour at the start of flow ramping up and after a flow peak compared to stable base flow. At the start of ramping up the flow, grayling made less position changes (movements >1 m) than at stable base flow and after a flow peak. In the unrestored habitat, the proportion of time grayling spent actively swimming was lower before a flow peak than it was both at the start of ramping up the flow and after the peak, an effect not found in the restored habitat. In addition, we compared two static flows, and habitat restoration mediated their effect on distance to nearest neighbour. Grayling in the restored habitat were positioned closer to each other in the low (~10 cm s−1) than in the intermediate static flow (~40 cm s−1), whereas in the unrestored habitat, grayling showed the opposite pattern. Moreover, grayling reduced their number of position changes in the intermediate static flow, which was reflected by a reduction in active swimming. Stomach analysis after the trials revealed that foraging success was higher in variable than in the stable flow treatment. These results show that flow magnitude, flow changes and instream structure play important roles in the behaviour of stream fishes.”

The paper is available open access on the journal website.

Johan Watz (left), together with Eirini Renata Karathanou and Antonis Apostolos Brouziotis.

The paper led by scientists of the Department of Engineering and Chemical Sciences, Karlstad University, was co-authored by Lutz Eckstein, professor of biology at the Department of Environmental and Life Sciences.

He writes about their work:

“Sweden has about 28 million hectares of forests, and pine trees constitute 40% of the total standing volume. Since the country is the world’s second largest exporter of pulp, paper and wood products, a total of 400 million containerized tree seedlings are produced by Swedish forest nurseries to restock forests each year. However, intensive annual forest harvests remove essential soil nutrients, which may cause problems for forest productivity.

In Sweden, container-grown seedlings are dominantly produced in peat and peat-based growth media. Peat-based substrates have many advantages such as long-term drainage ability, good aeration for tree seedling roots, good fertilizer absorbance and release capability. However, peat-based media are considered non-sustainable as their extraction have adverse environmental impacts. Therefore, sustainable approaches towards forest production and plantation management are urgently needed.

Therefore, the aim of this paper was to study the effects of hydrochar, derived from paper mill biosludge, on growth, quality, mycorrhizal associations and nutrient/heavy metal uptake of pine tree seedlings. We analyzed whether effects varied significantly between hydrochar forms (powder or pellets) or hydrochar proportions mixed with peat (10% or 20% hydrochar v/v). The effects of hydrochar addition on pine tree seedling was evaluated under three fertilization regimes (no fertilizer, 50% fertilizer and 100% fertilizer). We hypothesized that the growth, quality and mycorrhizal colonization of pine tree seedlings grown in substrate mixed with hydrochar would improve. We also expected pine tree seedlings grown with hydrochar to require less fertilizer to achieve similar or higher growth, mycorrhizal colonization and associated nutrient uptake relative to seedlings grown without hydrochar but with optimum rates of fertilizer (100% fertilizer). To our knowledge, this current study is the first paper to explore the potentials of hydrochar powder and pellets for being used as a growing media component in production of containerized pine tree seedlings.

Application of hydrochar had positive or neutral effects on shoot biomass and stem diameter compared with control seedlings (without hydrochar) under tested fertilizer levels. Analysis of the natural logarithmic response ratios (LnRR) of quality index and nutrient and heavy metal uptake revealed that application of 20% (v/v) hydrochar powder or pellet with 50% fertilizer resulted in same quality pine seedlings with similar heavy metal (Cu, Ni, Pb, Zn and Cr) and nutrient (P, K, Ca and Mg) contents as untreated seedlings supplied with 100% fertilizer. Colonization percentage by ectomycorrhizae significantly increased when either forms of hydrochar were applied at a rate of 20% under unfertilized condition. The results of this study implied that application of proper rates of hydrochar from biosludge with adjusted levels of liquid fertilizer may reduce fertilizer requirements in pine nurseries.”

Read the paper for free here! 

Johan Watz, Assistant Professor at Karlstad University, recently published two papers on juvenile salmonid ecology:


Temperature‐dependent competition between juvenile salmonids in small streams

By Johan Watz, Yasuhiko Otsuki, Kenta Nagatsuka, Koh Hasegawa & Itsuro Koizumi, published in the journal Freshwater Biology.

In the abstract, the authors write:

Johan Watz, doing field work during his PostDoc in Japan.

1) Biotic interactions affect species distributions, and environmental factors that influence these interactions can play a key role when range shifts in response to environmental change are modelled.

2) In a field experiment using enclosures, we studied the effects of the thermal habitat on intra‐ versus inter‐specific competition of juvenile Dolly Varden Salvelinus malma and white‐spotted charr Salvelinus leucomaenis, as measured by differences in specific growth rates during summer in allopatric and sympatric treatments. Previous laboratory experiments have shown mixed results regarding the importance of temperature‐dependent competitive abilities as a main driver for spatial segregation in stream fishes, and no study so far has confirmed its existence in natural streams.

3) Under natural conditions in areas where the two species occur in sympatry, Dolly Varden dominate spring‐fed tributaries (cold, stable thermal regime), whereas both species often coexist in non‐spring‐fed tributaries (warm, unstable thermal regime). Enclosures (charr density = 6 per m2) were placed in non‐spring‐fed (10–14°C) and spring‐fed (7–8°C) tributaries.

A forest stream on Hokkaido, northern Japan.

4) In enclosures placed in non‐spring‐fed tributaries, Dolly Varden grew 0.81% per day in allopatry and had negative growth (−0.33% per day) in sympatry, whereas growth rates were similar in allopatry and sympatry in spring‐fed tributaries (0.68 and 0.58% per day). White‐spotted charr grew better in sympatry than in allopatry in both thermal habitats. In non‐spring‐fed tributaries, they grew 0.17 and 0.79% per day and in spring‐fed tributaries 0.46 and 0.75% per day in allopatry and sympatry, respectively.

5) The negative effect of inter‐specific competition from white‐spotted charr on Dolly Varden thus depended on the thermal habitat. However, there was no strong evidence of a temperature‐dependent effect of intra‐ and inter‐specific competition on white‐spotted charr growth.

6) Multiple factors may shape species distribution patterns, and we show that temperature may mediate competitive outcomes and thus coexistence in stream fish. These effects of temperature will be important to incorporate into mechanistic and dynamic species distribution models.


Read more about the Koizumi lab at Hokkaido University (where Johan did his PostDoc) on their website!


Structural complexity in the hatchery rearing environment affects activity, resting metabolic rate and post‐release behaviour in brown trout Salmo trutta

By Johan Watz, published in the Journal of Fish Biology

In the abstract, Johan writes:

The effects of structural enrichment in the hatchery rearing environment of brown trout Salmo trutta was linked to post‐release performance. Enrichment resulted in reduced swimming activity scored in an open field test and reduced movement in a natural river after release. Also, enrichment increased resting metabolic rates, which correlated positively with overwinter growth.


Contact the author to access the papers.


The left photographs show Dolly Varden (a) and white‐spotted charr (b). The right photographs show enclosures in a non‐spring‐fed (c) and a spring‐fed (d) tributary.


The structurally enriched (left) and barren (right) tanks used in the study on how structural complexity in the hatchery environment affects juvenile brown trout.


River Rottnan in winter.

Yves P. Klinger, Sarah Harvolk-Schöning, Lutz Eckstein, Wiebke Hansen, Annette Otte and Kristin Ludewig recently published the paper ”Applying landscape structure analysis to assess the spatio-temporal distribution of an invasive legume in the Rhön UNESCO Biosphere Reserve” in the journal Biological Invasions.

Lutz Eckstein, Professor in Biology at Karlstad University, writes about their work:

The legume Lupinus polyphyllus. Photo by Lutz Eckstein

“We applied a combination of aerial mapping and GIS-based landscape analysis to study the invasion of the legume, Lupinus polyphyllus, in the Rhön UNESCO Biosphere Region as a case. We assessed the changes in lupine distribution between 1998 and 2016 in a strictly protected part of the Biosphere Region by means of landscape structure analysis. The area invaded by L. polyphyllus doubled from 1998 to 2016. The number of lupine stands decreased by 25%, but average stand size increased by 300%. In 2016, large and well-connected mesic grasslands that were situated close to roads were more heavily invaded than small and remote wet grasslands. Our results show that landscape composition plays an important role for the spread of invasive plants. Specifically, invasive stand characteristics, such as stand size, form, and connectivity, are crucial for driving the invasion process. Therefore, in addition to landscape composition, invasive stand characteristics should be included in the planning of conservation measures. Overall, aerial mapping combined with landscape analysis provides a cost-effective and practical tool for landscape managers to prioritize invasive control measures.”

Access the paper here, or contact any of the authors.

Åsa Enefalk, Ari Huusko, Pauliina Louhi and Eva Bergman recently published the paper “Fine stream wood decreases growth of juvenile brown trout (Salmo trutta L.)” in the journal Environmental Biology of Fishes. In the abstract, the authors write:

A juvenile brown trout (Salmo trutta) hiding in fine stream wood.

“In this study, the growth rate, gut fullness, diet composition and spatial distribution of brown trout was compared between artificial channels with and without fine wood (FW). Access to FW resulted in significantly lower brown trout growth rates over the study period from late summer to early winter as water temperatures declined from 17 °C to 1 °C. Access to FW resulted in minor differences in occurrence of the most common taxa found in brown trout diets, except for chironomid larvae which were found in c. 60% of the brown trout guts from control treatments but in only 30% of the guts from FW treatments in early winter. Diet consisted primarily of case-bearing and free-living Trichoptera larvae, Asellus, chironomid and Ephemeroptera larvae. Brown trout gut fullness was not significantly affected by access to FW bundles. Brown trout aggregated among FW but were more evenly distributed in channels lacking it. Our results suggest that juvenile brown trout use FW as a shelter at a wide range of water temperatures, and that this behaviour may result in reduced growth rates during their first fall and the onset of their first winter. We also show that prey availability and the composition of brown trout diet changes from late summer to early winter and that FW has a small but significant effect on brown trout diet composition.”

Read the paper here, or contact any of the authors.

Dirk Hattermann, Markus Bernhardt-Römermann, Annette Otte  and Lutz Eckstein recently published the paper “Geese are overlooked dispersal vectors for vascular plants in archipelago environments” in Journal of Vegetation Science.


In the abstract, the authors write:



We addressed the importance of gut‐mediated dispersal by Greylag Goose for vascular plants in archipelago environments and asked:

(i) What proportion of the local species pool is dispersed by geese?.

(ii) Which plant traits characterize species dispersed by geese?.

(iii) Which plant communities are likely to benefit from endozoochory by geese?.



Three Swedish Baltic archipelagos.



Goose droppings were collected on 45 islands. Plants germinating from the droppings represent the endozoochorous species pool (ESP). On 108 islands, the presence of vascular plants was recorded in each habitat. These species represent the island species pool (ISP). Differences in functional traits between ESP and ISP were expressed as effect sizes and tested using meta‐regressions. Using indicator species analyses and indicator species for managed semi‐natural grasslands, we identified the primary habitats of the ESP.



Geese dispersed viable diaspores of 97 plant species, which represents 22% of the ISP. Most ESP species were typical for small islands. Geese dispersed a higher proportion of graminoids and less woody plants, higher proportions of chamaephytes and therophytes and less phanerophytes; annuals and bi‐annuals were significantly overrepresented. One average, seed volume of the ESP was 95 % smaller than that of the ISP. About 51% of all ESP species were dispersed in at least two archipelagos. Geese showed a bias towards species of rocky shore habitats.



Geese potentially disperse large amounts of diaspores of many terrestrial island plant species. Through their feeding behaviour, geese select species with certain suites of traits from the regional species pool. Plant dispersal by geese may benefit plants species of rocky shores, but species of formerly managed semi‐natural grasslands may also find refuge sites on epilittoral shores after goose‐mediated dispersal. The relative importance of geese as dispersal vectors may increase under ongoing land‐use changes and cessation of grazing networks.”


Access the paper here, or contact any of the authors.

Two papers in Animal Conservation

Posted by Karl Filipsson | Papers

Two papers from NRRV were recently published in the journal Animal Conservation. The first paper presents a field study on how sedimentation affects brown trout (Salmo trutta) fry emergence in relation to freshwater pearl mussel (Margaritifera margaritifera) recruitment. The second paper presents a combined field and laboratory study on passage solutions for upstream-migrating eels (Anguilla anguilla).


Sedimentation affects emergence rate of host fish fry in unionoid mussel streams

Martin Österling


In the abstract, the author writes:

Freshwater pearl mussel, Margaritifera margaritifera

“Free-living, sympatric sedentary life stages of hosts and parasites are often adapted to similar environmental conditions. When the environment where these life stages occur is disturbed, both species can decline, causing strong negative effects on the parasitic species. For the highly threatened unionoid mussels with their larval parasitic life stage on fish, habitat degradation may simultaneously affect the conditions for the sedentary host fish eggs and the juvenile mussels in the sediment. This study provides novel information on the effect of sedimentation on the emergence rate of yolk sac fry, and its relation to mussel recruitment in two drainage basins, and is exemplified by the brown trout Salmo trutta, host fish for the threatened freshwater pearl mussel Margaritifera margaritifera. The results imply that turbidity and sedimentation can reduce the survival of trout eggs and yolk sac fry emergence rate regardless of trout strain and drainage basin. The results further suggest that low yolk sac fry emergence rates reduce the potential for mussel infestation and recruitment. The results indicate a year round negative effect of sedimentation, having strong and combined direct and indirect effects on juvenile mussel recruitment. Conservation measures that reduce anthropogenic sediment transportation into streams are a key factor for the conservation of mussels and their host fish.”

Access the paper here, or contact the author.


Climbing the ladder: an evaluation of three different anguillid eel climbing substrata and placement of upstream passage solutions at migration barriers

Johan Watz, Anders Nilsson, Erik Degerman, Carl Tamario and Olle Calles


European eel, Anguilla anguilla. Photo: Jörgen Wiklund

In the abstract, the authors write:

“Conservation programmes for endangered, long-lived and migratory species often have to target multiple life stages. The bottlenecks associated with the survival of juvenile anguillid eels migrating into inland waters, the survival and growth of the freshwater life stage, as well as the recruitment and survival of silver eels, migrating back to the ocean to spawn, must be resolved. In this study, we focus on the efficiency of passage solutions for upstream-migrating juveniles. Such solutions can consist of inclined ramps lined with wetted climbing substrata. We evaluated different commonly used substrata in a controlled experiment, recorded eel behaviour at the entrance of the ramp with infrared videography and validated the experimental results at a hydropower dam, where we also investigated the effects of ramp placement on performance. In the experiment on eel substratum selection, 40% of the eels passed in lanes with studded substratum, whereas only 21 and 5% passed using open weave and bristle substrata respectively. Video analysis revealed that the studded substratum attracted more approaches and initiated climbs than the other substrata, but once a climb had been initiated, passage success rates did not differ between substrata. Eels using the studded substratum climbed 26% faster than those using the bristle substratum and almost four times as fast as those climbing in the open weave. The superior performance of the studded substratum was supported by data from the field validation. Moreover, ramps positioned by the bank with low water velocities caught the most eels, but proximity to the dam had no effect on performance. To strengthen the European eel population, more juveniles need to reach their freshwater feeding grounds. A critical step to achieve this increase is to equip upstream passage solutions with suitable substrata and to optimize ramp placement at migration obstacles.”

Access the paper here, or contact any of the authors.

Viola elatior and the two habitats used in the study: early successional floodplain meadows and late successional alluvian woodlands. Photos: Benjamin Schulz.

Benjamin Schulz, Walter Durka, Jiří Danihelka and Lutz Eckstein recently published the research paper “Differential role of a persistent seed bank for genetic variation in early vs. late successional stages” in the scientific journal PLOS ONE. In the abstract, the authors write:

“Persistent seed banks are predicted to have an important impact on population genetic processes by increasing effective population size and storing past genetic diversity. Accordingly, persistent seed banks may buffer genetic effects of disturbance, fragmentation and/or selection. However, empirical studies surveying the relationship between aboveground and seed bank genetics under changing environments are scarce. Here, we compared genetic variation of aboveground and seed bank cohorts in 15 populations of the partially cleistogamous Viola elatior in two contrasting early and late successional habitats characterized by strong differences in light-availability and declining population size. Using AFLP markers, we found significantly higher aboveground than seed bank genetic diversity in early successional meadow but not in late successional woodland habitats. Moreover, individually, three of eight woodland populations even showed higher seed bank than aboveground diversity. Genetic differentiation among populations was very strong (фST = 0.8), but overall no significant differentiation could be detected between above ground and seed bank cohorts. Small scale spatial genetic structure was generally pronounced but was much stronger in meadow (Sp-statistic: aboveground: 0.60, seed bank: 0.32) than in woodland habitats (aboveground: 0.11; seed bank: 0.03). Our findings indicate that relative seed bank diversity (i.e. compared to aboveground diversity) increases with ongoing succession and despite decreasing population size. As corroborated by markedly lower small-scale genetic structure in late successional habitats, we suggest that the observed changes in relative seed bank diversity are driven by an increase of outcrossing rates. Persistent seed banks in Viola elatior hence will counteract effects of drift and selection, and assure a higher chance for the species’ long term persistence, particularly maintaining genetic variation in declining populations of late successional habitats and thus enhancing success rates of population recovery after disturbance events.”

Read the paper here!