Anne Ojala

Eddy covariance measurements of carbon exchange and latent and sensible heat fluxes over a boreal lake for a full open‐water period

was >0 W/m 2 at night and <0 W/m 2 in daytime. The latent heat flux dominated clearly over H in spring and summer; that is, the Bowen ratio was less than 1. Highermoment turbulence statistics proved to be efficient in detection of frequent nonstationary situations. Applying the statistical criteria for CO2 concentration and vertical wind speed, averaging over a 5-min period and selecting only the wind direction with longest fetch, we could obtain lake-representative CO2 fluxes. Footprint analysis based on a closure model revealed that the source areas were relatively short because of the presence of turbulence generated by the surrounding forest, compared to a larger lake with an extended smooth surface. We observed a net CO2 source of 0.2–0.4 mmol m � 2 s � 1 excluding July, when the flux was closer to zero. The results are consistent with the gradient method, based on more infrequent sampling, and both methods gave the same average flux, 0.2 mmol m � 2 s � 1 , over the whole open-water period.

Effects of cooling and internal wave motions on gas transfer coefficients in a boreal lake

Lakes and other inland waters contribute significantly to regional and global carbon budgets. Emissions from lakes are often computed as the product of a gas transfer coefficient, k 600 , and the difference in concentration across the diffusive boundary layer at the air–water interface. Eddy covariance (EC) techniques are increasingly being used in lacustrine gas flux studies and tend to report higher values for derived k 600 than other approaches. Using results from an EC study of a small, boreal lake, we modelled k 600 using a boundary-layer approach that included wind shear and cooling. During stratification, fluxes estimated by EC occasionally were higher than those obtained by our models. The high fluxes co-occurred with winds strong enough to induce deflections of the thermocline. We attribute the higher measured fluxes to upwelling-induced spatial variability in surface concentrations of CO2 within the EC footprint. We modelled the increased gas concentrations due to the upwelling and corrected our k 600 values using these higher CO2 concentrations. This approach led to greater congruence between measured and modelled k values during the stratified period. k 600 has a well-resolved and ~cubic relationship with wind speed when the water column is unstratified and the dissolved gases well mixed. During stratification and using the corrected k 600 , the same pattern is evident at higher winds, but k 600 has a median value of ~7 cm h−1 when winds are less than 6 m s−1, similar to observations in recent oceanographic studies. Our models for k 600 provide estimates of gas evasion at least 200% higher than earlier wind-based models. Our improved k 600 estimates emphasize the need for integrating within lake physics into models of greenhouse gas evasion.

Growth of Daphnia longispina L. in a polyhumic lake under various availabilities of algal, bacterial and detrital food

The availability and importance of food sources for growth of Daphnia longispina L. from a highly coloured fishless lake with anoxic hypolimnion were assessed by combining in situ and laboratory experiments. In in situ experiments populations were enclosed in tubes with natural temperature stratification and with or without anoxic hypolimnion. In the laboratory experiments the importance of food source (littoral zone vs pelagic epilimnion) was assessed by enclosing moss thalli and a natural zooplankton population in a large-scale flow-through system supplying food for experimental Daphnia. Growth of juveniles of Daphnia in epilimnetic water was determined in batch culture experiments and the importance of increasing concentrations of bacteria and algae for their growth and development was investigated with a small-scale flow-through system. Access to the anoxic hypolimnion enhanced the growth of Daphnia by 23–24%. Growth rates in the tubes with anoxic hypolimnion were 0.36 and 0.16 d−1 in July and August respectively. In tubes without anoxia the corresponding values were 0.29 and 0.13. In batch-cultures the highest growth rate determined was 0.16 and the overall rates were lower than in in situ experiments. In batch culture Daphnia was able to grow in darkness for 10 days with a rate of 0.16. In the large-scale flow-through system Daphnia population fed with littoral water reproduced well despite the low concentration of algae and increased its number by a factor of c. 32 in 10 days. However, the animals were small and net production of Daphnia population thus lower under the littoral influence than in the control treatment. Population could survive and grew slowly on pelagial water processed by a natural zooplankton community and with very little algae left. It is thus possible that bacteria serve as a ‘life-support system’ enabling the population survival over periods of algal shortage. Small-scale flow-through experiments revealed that Daphnia longispina is able to mature and reproduce on a bacterial diet if the food concentration is high enough and Daphnia on bacterial food could achieve growth rates similar to those on an algal diet. The threshold food level for Daphnia longispina was estimated to be c. 18.5 μg C 1−1. Detrital material is of limited value in nutrition of Daphnia even in a lake where more than 75% of carbon is bound in particulate detritus.

Carbon dioxide and energy fluxes over a small boreal lake in Southern Finland

Dynamics of carbon dioxide and energy exchange over a small boreal lake were investigated. Flux measurements have been carried out by the eddy covariance technique during two open-water periods (June–October) at Lake Kuivajarvi in Finland. Sensible heat (H) flux peaked in the early morning, and upward sensible heat flux at night results in unstable stratification over the lake. Minimum H was measured in the late afternoon, often resulting in adiabatic conditions or slightly stable stratification over the lake. The latent heat flux (LE) showed a different pattern, peaking in the afternoon and having a minimum at night. High correlation (r2 = 0.75) between H and water-air temperature difference multiplied by wind speed (U) was found, while LE strongly correlated with the water vapor pressure deficit multiplied by U (r2 = 0.78). Monthly average values of energy balance closure ranged between 70 and 99%. The lake acted as net source of carbon dioxide, and the measured flux (FCO2) averaged over the two open-water periods (0.7 µmol m−2 s−1) was up to 3 times higher than those reported in other studies. Furthermore, it was found that during period of high wind speed (>3 m s−1) shear-induced water turbulence controls the water-air gas transfer efficiency. However, under calm nighttime conditions, FCO2 was poorly correlated with the difference between the water and the equilibrium CO2 concentrations multiplied by U. Nighttime cooling of surface water enhances the gas transfer efficiency through buoyancy-driven turbulent mixing, and simple wind speed-based transfer velocity models strongly underestimate FCO2.

Effects of water clarity on lake stratification and lake‐atmosphere heat exchange

Recent progress of including lake subroutines in numerical weather prediction (NWP) models has led to more accurate forecasts. In lake models, one essential parameter is water clarity, parameterized via the light extinction coefficient, Kd, for which a global constant value is usually used. We used direct eddy covariance fluxes and basic meteorological measurements coupled with lake water temperature and clarity measurements from a boreal lake to estimate the performance of two lake models, LAKE and FLake. These models represent two 1D modeling frameworks broadly used in NWP. The results show that the lake models are very sensitive to changes in Kd when it is lower than 0.5 m−1. The progress of thermal stratification depended strongly on Kd. In dark water simulations the mixed layer was shallower, longwave and turbulent heat losses higher and therefore the average water column temperatures lower than in clear water simulations. Thus, changes in water clarity can also affect the onset of ice cover. The more complex LAKE modeled the seasonal thermocline deepening whereas it remained virtually constant during summer in the FLake model. Both models overestimated the surface water temperatures by about 1°C and latent heat flux by >30%, but the variation in heat storage and sensible heat flux were adequately simulated. Our results suggest that, at least for humic lakes, a lake-specific, but not time-depending, constant value for Kd can be used and that a global mapping of Kd would be most beneficial in regions with relatively clear lakes, e.g. in lakes at high altitudes.

Quality of detritus impacts on spatial variation of methane emissions from littoral sediment of a boreal lake

Spatial variation in CH 4 emissions in relation to net primary production (NPP) of vegetation was studied in a reed (Phragmites australis) -dominated littoral zone of a boreal meso-eutrophic lake during three growing seasons. In a healthy reed stand close to the shoreline, the proportion of CH 4 released to the atmosphere varied between 4 % and 5 % of NPP of P. australis. In a continuously flooded sparse reed stand in the middle infralittoral zone, this proportion was higher; plant-mediated emission was 18-24% of the NPP by P. australis, and the proportion of total emission (including ebullition) related to the NPP of all vegetation (including algae and Lemma trisulca) was 15-20%. In a batch experiment in the laboratory, detritus from L. trisulca and littoral algae maintained significantly higher CH 4 production than detritus from P. australis. The results support the hypothesis that the high spatial and temporal variability in CH 4 emissions in the littoral zone of the study lake was influenced by the quality and distribution of detritus in the anaerobic sediment.

Precipitation and net ecosystem exchange are the most important drivers of DOC flux in upland boreal catchments

According to recent studies, dissolved organic carbon (DOC) concentrations in rivers throughout the boreal zone are increasing. However, the mechanistic explanation of this phenomenon is not yet well known. We studied how the short and long-term changes in precipitation, soil temperature, soil water content, and net ecosystem exchange (NEE) are reflected to DOC concentrations and runoff DOC fluxes in two small forested upland catchments in Southern Finland. We used continuous eddy covariance measurements above the forest and runoff flow measurements from the catchment areas conducted over a 15 year long time period to study the correlation between NEE, gross photosynthetic production, total ecosystem respiration, litter production, and runoff DOC. In addition, we looked for the most important environmental variables in explaining the interannual changes in runoff DOC by using multiple linear regression. Finally, we studied the temporal connection between runoff DOC concentrations, precipitation, soil water content, and NEE by using wavelet coherence analysis technique. Our results indicate that the DOC concentrations have increased over the last 15 years. The DOC flux was to a large extent determined by the amount of precipitation, but the previous years NEE and litter production had also a small but significant effect on runoff DOC fluxes.

An urban boreal lake basin as a source of CO2 and CH4

Up to now, carbon gas fluxes from urban lakes in the boreal zone have seldom been studied. In summer 2005 we investigated fluxes from an urban boreal lake basin in southern Finland with long history of eutrophication and anoxia. Hypolimnetic CO₂ and CH₄ concentrations were high compared to other boreal lakes. During the open-water period, the lake basin acted as a source of CO₂ and CH₄ with fluxes of 2.10 mol m(-2 )and 0.04 mol m(-2), respectively. Despite the high oxidation rate (83%), CH₄ flux was higher than in other lakes and CH₄ contributed 33% [ corrected] to Global Warming Potential. The ratio of carbon emission to accumulation was 4, i.e. emissions were an important route for carbon departure but less so than in rural lakes. Since the lake oxygen conditions affected nutrient availability, there was a positive feedback from hypolimnion to carbon uptake, which was reflected in gas concentrations.

Robust parameters confirm predominance of heterotrophic processes in the plankton of a highly humic pond

The conclusions about the role of allochthonous organic matter in the food chains of inland waters are still often conflicting. Here we studied whether the results of specific experiments made in a small pond with very high concentration of allochthonous organic matter could be verified by simple, but robust, basic environmental parameters measured in the field. In summer primary production of phytoplankton could explain only ca. 20% of epilimnetic respiration and <10% of the nutritional requirements of zooplankton. These results agree with those of earlier experimental approaches and unequivocally suggest the major role played by allochthonous nutrition for the plankton in this pond. The roles played by photochemical degradation and anaerobic processes in the transformation of humic compounds available to epilimnetic food web in particular deserve further elucidation.

Combined Effects of Turbulence and Different Predation Regimes on Zooplankton in Highly Colored Water—Implications for Environmental Change in Lakes

In aquatic ecosystems, predation is affected both by turbulence and visibility, but the combined effects are poorly known. Both factors are changing in lakes in the Northern Hemisphere; the average levels of turbulence are predicted to increase due to increasing wind activities, while water transparency is decreasing, e.g., due to variations in precipitation, and sediment resuspension. We explored experimentally how turbulence influenced the effects of planktivorous fish and invertebrate predators on zooplankton when it was combined with low visibility caused by high levels of water color. The study was conducted as a factorial design in 24 outdoor ponds, using the natural zooplankton community as a prey population. Perch and roach were used as vertebrate predators and Chaoborus flavicans larvae as invertebrate predators. In addition to calm conditions, the turbulent dissipation rate used in the experiments was 10−6 m2 s−3, and the water color was 140 mg Pt L−1. The results demonstrated that in a system dominated by invertebrates, predation pressure on cladocerans increased considerably under intermediate turbulence. Under calm conditions, chaoborids caused only a minor reduction in the crustacean biomass. The effect of fish predation on cladocerans was slightly reduced by turbulence, while predation on cyclopoids was strongly enhanced. Surprisingly, under turbulent conditions fish reduced cyclopoid biomass, whereas in calm water it increased in the presence of fish. We thus concluded that turbulence affects fish selectivity. The results suggested that in dystrophic invertebrate-dominated lakes, turbulence may severely affect the abundance of cladocerans. In fish-dominated dystrophic lakes, on the other hand, turbulence-induced changes in planktivory may considerably affect copepods instead of cladocerans. In lakes inhabited by both invertebrates and fish, the response of top-down regulation to turbulence resembles that in fish-dominated systems, due to intraguild predation. The changes in planktivorous predation induced by abiotic factors may possibly cascade to primary producers.