Pär Byström

Light limitation of nutrient-poor lake ecosystems.

Productivity denotes the rate of biomass synthesis in ecosystems and is a fundamental characteristic that frames ecosystem function and management. Limitation of productivity by nutrient availability is an established paradigm for lake ecosystems. Here, we assess the relevance of this paradigm for a majority of the world’s small, nutrient-poor lakes, with different concentrations of coloured organic matter. By comparing small unproductive lakes along a water colour gradient, we show that coloured terrestrial organic matter controls the key process for new biomass synthesis (the benthic primary production) through its effects on light attenuation. We also show that this translates into effects on production and biomass of higher trophic levels (benthic invertebrates and fish). These results are inconsistent with the idea that nutrient supply primarily controls lake productivity, and we propose that a large share of the world’s unproductive lakes, within natural variations of organic carbon and nutrient input, are limited by light and not by nutrients. We anticipate that our result will have implications for understanding lake ecosystem function and responses to environmental change. Catchment export of coloured organic matter is sensitive to short-term natural variability and long-term, large-scale changes, driven by climate and different anthropogenic influences. Consequently, changes in terrestrial carbon cycling will have pronounced effects on most lake ecosystems by mediating changes in light climate and productivity of lakes.

CANNIBALISM AND COMPETITION IN EURASIAN PERCH: POPULATION DYNAMICS OF AN ONTOGENETIC OMNIVORE

In many cannibalistic populations, cannibals share resources with their victims, leading to a size-dependent mixture of cannibalistic and competitive interactions. We analyze the impacts of such interactions on the population dynamics of Eurasian perch (Percafluviatilis) by considering effects of intercohort competition, habitat heterogeneity, habitat selection, and energy gain made by cannibals. Over a six-year period, we measured mortality and recruitment patterns, individual growth, body condition, resource levels, diets, and habitat use as functions of density for an allopatric perch population in a low-productivity lake. During the course of the study, two major die-offs took place, selectively affecting larger cannibalistic individuals, followed by several years of successful recruitment of young fish. Habitat use of perch ?2 yr old was density dependent, and these fish used only the inshore region at low densities. The appearance of young fish followed the die-offs of cannibalistic perch and their subsequent absence from the offshore area, both of which decreased cannibalism on pelagic recruits. Whereas die-offs of larger perch could not be related to competition with young-of-the-year (YOY) perch, evidence for a competitive impact of YOY perch on Age-I perch was present due to substantial food overlap. A strong depression in pelagic zooplankton was observed during summer in years with strong re- cruitment, which resulted in reduced consumption of zooplankton, slow growth, and reduced body condition in Age-1 perch and suggested high mortality of Age-1 perch in autumn. Age-I perch did not appear to profit substantially from cannibalism on YOY perch because of the short time period that they could efficiently prey on YOY perch. The few larger perch that survived the die-offs gained substantial energy from cannibalism in years with strong recruitments, which increased both growth rates and per capita fecundity. Size-dependent intercohort competition may have strong impacts on cannibal-victim in- teractions when victims share resources with cannibals. Furthermore, habitat heterogeneity, combined with habitat selection, may limit the extent to which cannibals have a stabilizing effect on population dynamics. Finally, the energy gained by cannibals may have important consequences on population dynamics as this energy is allocated into new recruits.

Density Dependent Growth and Size Specific Competitive Interactions in Young Fish

Body size is a major determinant affecting an individuals performance. In this study, four factors affecting an organisms competitive ability as related to body size: the attack rate, the maximum growth capacity, the metabolic demands and the size spectrum of available resources, were investigated for small stages of Eurasian perch (Percu fluviatilis) and roach (Rutilus rutilus). The size dependence of the attack rate on a 0.5-mm cladoceran zooplankton of larval and juvenile perch was estimated and compared with the results from similar experiments for roach. At equal body sizes, roach always had a higher attack rate than perch. In contrast the reverse was the case for maximum growth capacity. The maintenance requirements at the same size were higher for perch than for roach. Based on the above data we were able to gain a mechanistic understanding of the outcome of two field enclosure competition experiments, one between perch larvae and I-yr-old roach and one between perch larvae and roach larvae. In the first experiment, we found strong intraspecific density dependent growth in perch larvae, while the presence of I-yr-old roach had no effect on perch larvae growth. High perch densities had strong negative effects on the zooplankton resource. Due to the larger size of I-yr-old roach and consequently higher metabolic demands, I-yr-old roach showed a stronger negative response in growth to high perch larvae densities than perch despite roachs higher attack rate on zooplankton. In the second experiment, perch larvae were negatively affected by high densities of roach larvae and had a reduced growth at high intraspecific densities. In contrast, the growth of roach larvae was not affected by perch larvae or high intraspecific densities. This difference between species could be related to the simultaneous lower attack rate and higher growth capacity of perch, leading to a higher sensitivity in growth of perch than of roach to decreasing resource levels. Temporal variation in competition intensity was present in both experiments. This variation could be related to the foraging efficiencies and different growth capacities of the competitors and the species composition and size structure of the zooplankton resource. Our study points to the potential for both intra- and interspecific competitive interactions in fish larvae in freshwater systems. Our results also suggest that species specific differences in how foraging, growth and food processing capacities relate to body size are of vital importance for interactions in ecological communities.

Whole‐lake estimates of carbon flux through algae and bacteria in benthic and pelagic habitats of clear‐water lakes

This study quantified new biomass production of algae and bacteria in both benthic and pelagic habitats of clear-water lakes to contrast how carbon from the atmosphere and terrestrial sources regulates whole-lake metabolism. We studied four small unproductive lakes in subarctic northern Sweden during one summer season. The production of new biomass in both benthic and pelagic habitats was calculated as the sum of autotrophic production by algae and heterotrophic production by bacteria using allochthonous organic carbon (OC). Whole-lake production of new biomass was dominated by the benthic habitat (86% +/- 4% [mean +/- SD]) and by primary production (77% +/- 9%). Still, heterotrophic bacteria fueled by allochthonous OC constituted a significant portion of the new biomass production in both benthic (19% +/- 11%) and pelagic habitats (51% +/- 24%). In addition, overall net production (primary production minus respiration) was close to zero in the benthic habitats but highly negative (-163 +/- 81 mg C x m(-2) x d(-1)) in pelagic regions of all lakes. We conclude (1) that allochthonous OC supported a significant part of total production of new biomass in both pelagic and benthic habitats, (2) that benthic habitats dominated the whole-lake production of new biomass, and (3) that respiration and net CO2 production dominated the carbon flux of the pelagic habitats and biomass production dominated the benthic carbon flux. Taken together, these findings suggest that previous investigations have greatly underestimated the productivity of clear-water lakes when benthic autotrophic production and metabolism of allochthonous OC have not been measured.

Size-dependent foraging efficiency, cannibalism and zooplankton community structure

Abstract To examine size-dependent food web interactions in systems with cannibalism, we compared the abundances of zooplankton and phytoplankton over 2 years in four lakes with cannibalistic perch (Perca fluviatilis) of which two also supported the top predator pike (Esox lucius). The abundance of perch 2 years and older was lower in lakes with pike than in lakes with only perch. In contrast, the abundance of small perch (young-of-the-year and 1-year old) was lower in lakes with only perch suggesting that intense cannibalism reduced these size classes to low levels in lakes lacking pike. Functional response experiments with differently sized perch and zooplankton showed that the attack rate of small perch susceptible to cannibalism was much higher than that of large cannibalising perch. The optimal body size of perch with respect to attack rate was also lower for small zooplankton prey than for large zooplankton. The zooplankton communities in lakes with only perch were dominated by the relatively small species Ceriodaphnia quadrangula and Bosmina spp. and total zooplankton biomass was higher in these lakes than in lakes with both pike and perch. In contrast, the mean size of cladoceran zooplankton was largest in lakes with both pike and perch owing to a dominance of the large zooplankton species Holopedium gibberum in these lakes. We relate these patterns to (1) the low foraging efficiency of large perch on small zooplankton and (2) the low abundance of small zooplanktivorous perch (due to cannibalism) in lakes with only perch. The differences in zooplankton community structure also resulted in different seasonal dynamics of phytoplankton between lakes. Cannibalism introduces a vertical heterogeneity to food webs that causes consumer-resource dynamics that are not predictable from linear food chain models.

Resource limitation during early ontogeny: constraints induced by growth capacity in larval and juvenile fish

Abstract The presence of and mechanisms behind density-dependent growth and resource limitation in larval and juvenile stages of organisms with high mortality such as fish are much debated. We compare observed consumption and growth rates with maximum consumption and growth rates to study the extent of resource limitation in young-of-the-year (YOY) roach (Rutilus rutilus) and perch (Perca fluviatilis). Diet, habitat use, consumption rate and growth rate were measured under varying YOY fish densities over 2 years in four lakes. In the first year, YOY roach and perch were studied under allopatric conditions. Experimental addition of perch roe in the second year also allowed study of YOY of the two species under sympatric conditions in two of the lakes. The diet of YOY roach was dominated by cladoceran zooplankton and YOY roach habitat use was restricted to the shore region in both years. This restricted habitat use did not involve any cost in foraging gain in the first year as consumption and growth rates were very close to maximum rates. During the second year, when the two species coexisted, resources were limited in late season, more so in the littoral than in the pelagic habitat in one lake while the reverse was the case in the other lake. The diet of YOY perch was also dominated by zooplankton, and with increasing perch size the proportion of macroinvertebrate prey in the diet increased. After hatching, YOY perch first utilized the pelagic habitat restricting their habitat use to the shore after 1 to several weeks in the pelagic zone. During the larval period, perch were not resource limited whereas juvenile perch were resource limited in both years. The fact that YOY perch were more resource limited than YOY roach was related to the higher handling capacity and lower attack rate of perch relative to roach, rendering perch more prone to resource limitation. Estimates of resource limitation based on consumption rates and growth rates yielded similar results. This supports the adequacy of our approach to measure resource limitation and suggests that this method is useful for studying resource limitation in organisms with indeterminate growth. Our results support the view that density-dependent growth is rare in larval stages. We suggest that density-dependent growth was absent because larval perch and roach were feeding at maximum levels over a wide range of larvae densities.

Interactions among size-structured populations in a whole-lake experiment: size- and scale-dependent processes

Interactions in size-structured populations are characterized by a mixture of predatory and competitive interactions dependent on the size of the individual organism. We analyzed this mixture of size-structured interactions for a species constellation consisting of perch (Perca fluviatilis) and roach (Rutilus rutilus) in a replicated whole lake experiment over 4 yr. Roach are preyed upon by large perch, but at the same time compete with small perch for zooplankton. Predictions regarding the effects of roach on perch performance and resource dynamics in the whole-lake experiment were based on results from previous pond and enclosure experiments carried out over short (months) time periods. Variables measured in the whole-lake experiment included both individual level parameters (diet and growth of perch) as well as population level parameters (mortality, population numbers and size structures of perch, abundance and biomass of zooplankton and benthic macroinvertebrates). The population size of perch ≥ 2 yr old decreased to less than 10% of the pre-treatment levels in treatment lakes after the introduction of roach and remained low over the whole study period. The mortality in perch one-year-old and older increased with size. Larger perch had a higher growth decrease than smaller perch, but also the individual growth rate of young-of-the-year (YOY) perch was lower in treatment lakes than in control lakes. YOY perch were almost excluded during the winter and spring following the roach introduction. The few perch of this year class that survived had the following years a higher growth than the corresponding year class in control lakes. This could be related to an increased availability of predator-sensitive macroinvertebrates in treatment lakes. Perch in treatment lakes fed to a larger extent on macroinvertebrates and less on pelagic zooplankton than perch in control lakes. As expected, zooplankton abundances decreased in treatment lakes the year following the roach introduction, but thereafter zooplankton abundances were higher in treatment lakes due to the absence of perch recruitment in these lakes. No incidence of perch predation on roach was recorded during the study period. We attribute this to the spatial scale dependent escape ability and habitat use of vulnerable size classes of roach. YOY perch had a stronger impact on between-year variation in zooplankton than YOY roach and this can be related to the higher foraging (handling) capacity of the former and the restricted habitat use of the latter. Individual level parameters (diet, growth) had a higher statistical power than population level parameters (abundance, biomass, mortality), and small-scale experiments successfully predicted a higher proportion (81%) of the individual level parameters. However, the lower success in predicting population level responses (38%) was directly connected to a failure to predict individual level parameters (large perch diet and growth). Our results suggest that the small spatial scale was the major limitation on the predictive ability of the small-scale experiments.

Resource heterogeneity, diet shifts and intra-cohort competition: effects on size divergence in YOY fish

Most organisms exhibit a substantial size variation among individuals due to individual differences in experienced biotic and abiotic environmental conditions and because individuals undergo growth and development during most of their life time. One important issue in this context is how size variation within cohorts may develop over time. Here, we tested the hypothesis, in gape-limited animals such as fish, that size divergence among individuals within a cohort depends on the opportunity to undergo size-dependent diet shifts, by allowing initially larger individuals to make an early diet shift when the first resource becomes limiting. We used young-of-the-year perch (Perca fluviatilis) as our study organism. Competitive intensity and the opportunity to undergo a diet shift from zooplankton to macroinvertebrates affected both mean growth rates and the extent to which inter-individual variation in growth was manifested. As predicted, increased competition combined with the presence of both zooplankton and benthic macroinvertebrates increased the degree of size variation. However, size divergence was also observed among individuals when only the initial resource, zooplankton, was available. We argue that only non-exploitative interactions, such as dominance structures and social interactions, could have caused this latter pattern, as exploitative competition is expected to lead to size convergence due to the superior competitive ability of smaller individuals. Our results suggest that diet shifts are not a prerequisite for size divergence in animal cohorts, and that dominance and social interactions may have similar effects on size variation within cohorts. Finally, development of size variation is suggested to have strong implications for overall cohort performance.

Terrestrial organic matter input suppresses biomass production in lake ecosystems

Terrestrial ecosystems export large amounts of organic carbon (t-OC) but the net effect of this OC on the productivity of recipient aquatic ecosystems is largely unknown. In this study of boreal lakes, we show that the relative contribution of t-OC to individual top consumer (fish) biomass production, and to most of their potential prey organisms, increased with the concentration of dissolved organic carbon (DOC; dominated by t-OC sources) in water. However, the biomass and production of top consumers decreased with increasing concentration of DOC, despite their substantial use (up to 60%) of t-OC. Thus, the results suggest that although t-OC supports individual consumer growth in lakes to a large extent, t-OC input suppresses rather than subsidizes population biomass production.

Impacts of elevated terrestrial nutrient loads and temperature on pelagic food-web efficiency and fish production

Both temperature and terrestrial organic matter have strong impacts on aquatic food-web dynamics and production. Temperature affects vital rates of all organisms, and terrestrial organic matter can act both as an energy source for lower trophic levels, while simultaneously reducing light availability for autotrophic production. As climate change predictions for the Baltic Sea and elsewhere suggest increases in both terrestrial matter runoff and increases in temperature, we studied the effects on pelagic food-web dynamics and food-web efficiency in a plausible future scenario with respect to these abiotic variables in a large-scale mesocosm experiment. Total basal (phytoplankton plus bacterial) production was slightly reduced when only increasing temperatures, but was otherwise similar across all other treatments. Separate increases in nutrient loads and temperature decreased the ratio of autotrophic:heterotrophic production, but the combined treatment of elevated temperature and terrestrial nutrient loads increased both fish production and food-web efficiency. CDOM: Chl a ratios strongly indicated that terrestrial and not autotrophic carbon was the main energy source in these food webs and our results also showed that zooplankton biomass was positively correlated with increased bacterial production. Concomitantly, biomass of the dominant calanoid copepod Acartia sp. increased as an effect of increased temperature. As the combined effects of increased temperature and terrestrial organic nutrient loads were required to increase zooplankton abundance and fish production, conclusions about effects of climate change on food-web dynamics and fish production must be based on realistic combinations of several abiotic factors. Moreover, our results question established notions on the net inefficiency of heterotrophic carbon transfer to the top of the food web.