Kalevi Salonen

Perspectives in Winter Limnology: Closing the annual cycle of freezing lakes

Winter has traditionally been considered as an ecologically insignificant season and, together with technical difficulties, this has led winter limnology to lag behind summer limnology. Recently, rapidly expanding interest in climate warming has increased water research in winter. It has also become clear that neither winter conditions of lakes nor under-ice communities are as static as often supposed. Although interannual differences in water temperature are small, close to the maximum density temperature, they may have profound effect on under-ice hydrodynamics. Thus, stochastic variations in weather, particularly those preceding the time of freezing and ice melting, may have important consequences for hydrodynamics which then affect the distributions and conditions of microorganisms and probably further to higher trophic levels. Even fish distributions can be dictated by under-ice conditions and their activities as well as behavior can sometimes approach those in summer. Life in freshwater ice is one of the least studied aspects of winter limnology and recent studies suggest that a thorough evaluation is needed. Altogether there are strengthening signs that winter should be considered as an integral part in the functioning and dynamics of lakes affecting quantitative and qualitative characteristics of aquatic communities in summer. There are great prospects that more thorough understanding of the prevailing limnological conditions in winter will improve our understanding of lake ecosystems in their entirety, and there is no doubt that such an approach requires multidisciplinary and long- term studies at different spatial scales.

Phytoplankton in Lake Tanganyika : vertical and horizontal distribution of in vivo fluorescence

Determinations of chlorophyll a and in vivo fluorescence of photosynthetic pigments were used to study vertical and horizontal distribution of phytoplankton in Lake Tanganyika (East Africa). Blue excited fluorescence (IVFb) was an approximate predictor of chlorophyll a at different depths and locations. Green excited fluorescence (IVFg), which reflects phycoerythrin in cyanobacteria, explained chlorophyll a variation equally well, and in combination with IVFb the degree of explanation was improved to 87% (n = 90). Particularly during the shallow stratification in March–May, the maxima of chlorophyll a, IVFb and IVFg were located within the thermocline. Such distribution may have resulted from the high penetration of UV light, often accentuated by very shallow daytime thermal stratification, leading to inhibition of phytoplankton near the surface. Because the decrease of chlorophyll a specific IVFb was less striking towards the surface, the decrease of IVFb was not caused by light inhibition only. In October–November, epilimnetic IVFb and chlorophyll a values seemed to be consistently higher than in April–May and often showed remarkable patchiness. The sometimes very dense phytoplankton blooms (Anabaena sp., Cyanobacteria) observed in the central and southern parts of the lake, suggest that local upwelling or mixing events may be important for the development of phytoplankton in Lake Tanganyika.

The stoichiometry of particulate nutrients in Lake Tanganyika — implications for nutrient limitation of phytoplankton

We studied the potential nutrient limitation of phytoplankton by means of seston nutrient stoichiometry and nutrient enrichment bioassays in the epilimnion of Lake Tanganyika. In most cases, the particulate carbon to phosphorus (C:P) ratio was high and indicated moderate P deficiency, while the respective C:N ratio mainly suggested moderate N deficiency. The N:P ratios of seston indicated rather balanced N and P supply. In three two-day enrichment bioassays in April–May 1995, a combined addition of P, N and organic carbon (glucose) always increased primary production in comparison to untreated controls. Primary production also slightly increased after the addition of phosphate-P, while the additions of single ammonium-N and glucose had no effect. Although the measured turnover time of P was short and our few nutrient enrichment experiments suggested that P may be the most limiting single nutrient, the particulate nutrient ratios and the strong stimulation of primary production after the combined addition of P and N mostly suggest that in the upper epilimnion of Lake Tanganyika plankton experience a restricted, but approximately balanced nutrient supply.

Radiation transfer and heat budget during the ice season in Lake Pääjärvi, Finland

Lake Pääjärvi, a boreal Finnish lake, was investigated in winter for weather conditions, structure and thickness of ice and snow, solar radiation, and under-ice current and temperature. Heat budget of Lake Pääjärvi in January–March was governed by terrestrial radiation losses of 20–35xa0Wxa0m−2 recompensed by ice growth of 0.5–1.0xa0cmxa0day−1. In April, snow melted, albedo decreased from 0.8 to <0.1, and the mean ice melt rate was 1.5xa0cmxa0day−1. Internal melting and surface melting were about equal. The mean turbulent heat loss was small. The heat flux from the water to ice was about 5xa0Wxa0m−2 in winter, increasing to 12xa0Wxa0m−2 in the melting season. The light attenuation coefficient was 1.1xa0m−1 for the congelation ice (black ice) in winter, compared with 1.5xa0m−1 for the lake water, and it was up to 3xa0m−1 for candled congelation ice in spring, and about 10xa0m−1 for superimposed ice (white ice) and snow. Gas bubbles were the main factor that reduced the transparency of ice. The radiation penetrating the ice heated the water body causing convective currents and horizontal heat transfer. This increased the temperature of the water body to about 3°C before the ice break-up. After the snow had melted, the euphotic depth (the depth of 1% surface irradiance) was estimated as 2.0xa0m, only two-thirds that in summer.

Bacterioplankton in a small polyhumic lake with an anoxic hypolimnion

Bacterioplankton biomass and dark fixation of inorganic carbon were measured in the highly humic (water colour up to 550 mg Pt 1-1) and acidic lake, Mekkojarvi. Strong thermal and chemical stratification developed in the water column early in spring and led rapidly to anoxia in the hypolimnion, which extended to less than 1.0 m from the surface. In the epilimnion only small bacteria were abundant. In the anoxie zone both the abundance and the mean size of bacteria were considerably higher than in the epilimnion. These differences are thought to be the result of different grazing pressure from Zooplankton in the two zones. In late summer a high concentration of bacteriochlorophyll d in the upper hypolimnion indicated a high density of photo synthetic bacteria. Bacterial biomass was similar to that of phytoplankton in the epilimnion, but 23 times higher in the whole water column. In August, dark fixation of inorganic radiocarbon in the anaerobic zone was 51% of the total 14C-incorporation and the contribution of light fixation was only 5.4%. In the polyhumic Mekkojarvi, bacterioplankton was evidently a potentially significant carbon source for higher trophic levels, but bacterioplankton production could not be supported by phytoplankton alone. Allochthonous inputs of dissolved organic matter probably support most of the bacterial production.

Development of phytoplankton in Lake Pääjärvi (Finland) during under-ice convective mixing period

The development of winter phytoplankton communities was studied in both shallow and deep areas of Lake Pääjärvi, southern Finland, during the final 2xa0weeks of winter ice cover. Phytoplankton was mainly composed of diatoms, cryptophytes and chrysophytes. The diatoms Aulacoseira and Rhizosolenia were always uniformly distributed with depth, initially probably due to mixing induced by heat flux from the sediment and later due to thermal convection. Motile Rhodomonas cryptophytes and Chrysococcus chrysophytes were most abundant near the ice showing that, despite their small size, they were partly able to resist mixing by convection. Their ability to stay in more illuminated water layers was reflected in net rates of increase about an order of magnitude higher than those of diatoms in the middle of the lake. Given the low temperatures and convection, the observed net rates of increase of motile taxa were very high compared to growth rates reported in the literature. The gradual increase in light availability following melting of ice led to a consistent increase in the abundances of major phytoplankton taxa irrespective of deep convective circulation. It is suggested that those algae most abundant at the time of ice break have a competitive advantage in the following open water conditions when nutrients are abundant but deep water circulation limits light availability. The results emphasize that in lakes which cool below the maximum density of water before freezing, apparently small differences in temperature and light conditions can cause important changes in the circulation patterns that impact on phytoplankton development.

Spatial Distribution of Phytoplankton and Picocyanobacteria in Lake Tanganyika in March and April 1998

The spatial distribution of phytoplankton and picocyanobacteria was studied in Lake Tanganyika in relation to environmental factors. Sampling was conducted within three weeks during the wet season of 1998 when the water column stratification was most stable. The secchi depth varied between 11 and 16 m, except off the river Malagarasi (3 m). The depth of the euphotic zone was 33 to 56 m. Altogether, 218 phytoplankton taxa were identified. Their total biomass varied between 13 and 88 mg m m 3 fresh weight. Picocyanobacteria were present at very high densities (10 4 to 6 2 10 5 cells ml m 1 ). The influence of the river Malagarasi appeared to spread along the water surface because of the lower density of the river water. The phytoplankton species number and biomass were particularly high near the river mouth. Excluding that area, there were no statistically significant differences in the species composition among study sites. Nevertheless, differences in water column mixing, indicated by variable temperature gradients, seemed to affect the horizontal distribution of at least some phytoplankton species.

Comparison of thermal stratification, light attenuation and Chlorophyll-a dynamics between the ends of Lake Tanganyika

Thermal structure, light attenuation, and chlorophyll- a dynamics within the upper 100 m at the northern and southern ends of Lake Tanganyika were measured from August 1995-July 1996. Pronounced variability of thermal structure in time, depth and region were observed. During the dry windy season (June-September), the water column in the south was largely isothermal down to 100 m, while in the north stratification was detectable. Subsequently, in October, cessation of winds coincided with a reestablishment and strengthening of thermal stratification in the south, and a distinct rise of a weakened thermocline in the north was accompanied by an increase in epilimnetic concentrations of dissolved solids. The depth of photic zone was highly variable with the average position of 1% of incident photosynthetically active radiance found between 20 and 70 m, sometimes showing vertical shifts of 25 m a week. Weekly observations showed that concentrations of chlorophyll- a were rather similar at both ends of the lake. At both stations, periods of unstable or absent thermal stratification were accompanied by increased concentrations of chlorophyll- a , although in the north chlorophyll- a also peaked during periods of deep thermal stratification and low rates of mixing. Concentrations of chlorophyll- a did not show significant relationship with the available amount of underwater available photosynthetically active radiance (I m from 11-173 w Em -2 s -1 ). We suggest that in Lake Tanganyika chlorophyll- a maxima can occur whenever growth rates exceed spatial mixing rates, that is, if mixing is less than a critical turbulence. Our observations underscore the consequences of hydrodynamic processes on biological productivity either by securing internal nutrients or by controlling the timing and magnitude of phytoplankton biomass production.

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.

Size-Fractionated δ 15 N and δ 13 C Isotope Ratios Elucidate the Role of the Microbial Food Web in the Pelagial of Lake Tanganyika

Food web structure of the pelagic community in Lake Tanganyika was studied using the stable nitrogen and carbon isotopes 15 N and 13 C. Size-fractionated seston, zooplankton, shrimps, medusae and fish were sampled in the northern part of Lake Tanganyika. Picoplankton fractions as well as cyanobacteria-dominated nano/microplankton fractions had very low nitrogen isotope signatures typical for nitrogen-fixing organisms. Fractions containing mainly dead organic matter (and associated bacteria) or nano/microalgae (chlorophytes and diatoms) had i 15 N 2 to 4 higher. The low i 15 N signatures of small cyclopoids and shrimps suggest they are feeding on nitrogen-fixing cyanobacteria (picoplankton or larger forms), while the higher i 15 N signature of larger copepods suggest mixed feeding on large algae and small zooplankton and/or cyanobacteria. Medusae were slightly enriched in i 15 N relative to large copepods. Among fish, the signatures of Stolothrissa and small Lates stappersi suggested feeding on large copepods, while Limnothrissa and larger Lates were slightly more enriched, indicating partial piscivory. The enrichment of 13 C between the putative trophic levels (2 to 3) was higher, while that of 15 N (2 to 3) was lower, than usual in isotope studies. Our results indicate that picocyanobacteria and possibly also larger cyanobacteria are important producers in the pelagic food web of Tanganyika.