Nitrification and Denitrification in High-Strength Ammonium by <Emphasis Type="Italic">Alcaligenes faecalis</Emphasis>

Alcaligenes faecalis sp. No. 4, that has the ability of heterotrophic nitrification and aerobic denitrification in high-strength ammonium at about 1200 mg-N/l, converted about one-half of removed NH ₄⁺-N to intracellular nitrogen and nitrified only 3% of the removed NH₄⁺. From the nitrogen balance, 40–50% of removed NH₄⁺-N was estimated to be denitrified. Production of N₂ was confirmed by GC-MS and 90% of denitrified products was N₂. The maximum ammonium removal rate, 29 mg-N/l h and its denitrification rate in aerated batch experiments, were 5–40 times higher than those of other bacteria with the same ability.     

RESPONSE OF CATALASE ACTIVITY TO ENVIRONMENTAL STRESS IN THE FRESHWATER DINOFLAGELLATE PERIDINIUM GATUNENSE1

Catalase activity increased in Peridinium gatunense (formerly P. cinctum fa. westii) cells during the decline of the seasonal spring bloom period in Lake Kinneret. This was correlated with the low ambient total CO₂ concentration. The relationship was confirmed in laboratory experiments where maximum catalase activity occurred under an atmosphere composed of 30% O₂ and 0.003% CO₂. Conversely, high CO₂ concentrations inhibited catalase activity. The rise in catalase activity was not directly due to increasing environmental pH, as in vitro and in vivo measurements showed a characteristic broad pH curve with a constant activity from pH 6–10 for catalase. Photoinhibition of catalase occurred above 250 μmol photons · m^?2· s^?1. However, at high photoinactivating irradiances, photoinhibition was ameliorated under high pO₂/pCO₂. Such conditions prevail in the Kinneret at the end of the spring. We propose that the enhancement of photorespiration (under high pO₂/pCO₂) induces a temporary burst in catalase activity despite the progressively photoinhibitory conditions of early summer.     

Appearance and establishment of diazotrophic cyanobacteria in Lake Kinneret,Israel

1. We propose that the appearance and establishment of Nostocales (cyanobacteria) species of the genera Aphanizomenon and Cylindrospermopsis in the warm subtropical Lake Kinneret (Sea of Galilee, Israel) from 1994 was linked to changes in climate conditions and summer nitrogen (N) availability. 2. From 1994 to 2009, an increase in frequency of events of elevated water temperature (>29 °C) in summer, and to some extent a greater frequency of lower summer wind speed events, affected water turbulence and water column stratification, thus providing better physical conditions for the establishment of these populations. 3. In recent years, N‐depleted conditions in Lake Kinneret in early summer have promoted the development and domination of Nostocales that could gain an ecological advantage owing to their N₂‐fixing capability. 4. Nitrogen fixation rates coincided both with heterocyst abundance and with Nostocales biomass. The N supplied to the lake via nitrogen fixation ranged from negligible quantities when Nostocales represented only a minor component of the phytoplankton community to 123 tonnes when Cylindrospermopsis bloomed in 2005. This high N₂ fixation rate equals the average summer dissolved inorganic nitrogen load to the lake via the Jordan River.     

Heterotrophic nitrogen removal by <Emphasis Type="Italic">Providencia rettgeri</Emphasis> strain YL

Providencia rettgeri strain YL was found to be efficient in heterotrophic nitrogen removal under aerobic conditions. Maximum removal of NH₄ ⁺–N occurred under the conditions of pH 7 and supplemented with glucose as the carbon source. Inorganic ions such as Mg²⁺, Mn²⁺, and Zn²⁺ largely influenced the growth and nitrogen removal efficiency. A quantitative detection of nitrogen gas by gas chromatography was conducted to evaluate the nitrogen removal by strain YL. From the nitrogen balance during heterotrophic growth with 180 mg/l of NH₄ ⁺–N, 44.5% of NH₄ ⁺–N was in the form of N₂ and 49.7% was found in biomass, with only a trace amount of either nitrite or nitrate. The utilization of nitrite and nitrate during the ammonium removal process demonstrated that the nitrogen removal pathway by strain YL was heterotrophic nitrification-aerobic denitrification. A further enzyme assay of nitrate reductase and nitrite reductase activity under the aerobic condition confirmed this nitrogen removal pathway.     

Nitrous oxide in brackish Lake Nakaumi, Japan II: the role of nitrification and denitrification in N2O accumulation

In order to understand the role of nitrification and denitrification in the accumulation of nitrous oxide (N₂O) in the hypolimnetic water of brackish Lake Nakaumi, the effects of dissolved oxygen (DO) concentration on these activities were investigated by incubation experiments. N₂O was produced during the oxidation of NH₄ ⁺ to NO₂ ⁻ in nitrification and during the reduction of NO₃ ⁻ to N₂ in denitrification. N₂O-producing activity by nitrification (N₂O_N) increased markedly with decreasing concentrations of DO. Low DO (10%–30% saturation) induced high N₂O_N. In contrast to nitrification, N₂O-producing activity by denitrification (N₂O_D) decreased with decreasing concentrations of DO. Little N₂O was accumulated during denitrification under low-level conditions of DO (10%–30%), because of further reduction of N₂O to N₂. It can therefore be assumed that N₂O produced as the by-product of nitrification is concurrently reduced to N₂ by denitrification under low-DO conditions. This would result in no substantial accumulation of N₂O during active nitrification in the hypolimnetic water of Lake Nakaumi. Received: July 6, 2001 / Accepted: December 10, 2001     

Nitrogen removal in a wastewater treatment plant through biofilters: nitrous oxide emissions during nitrification and denitrification

In order to estimate N₂O emissions from immersed biofilters during nitrogen removal in tertiary treatments at urban wastewater treatment plants (WWTPs), a fixed culture from the WWTP of “Seine Centre” (Paris conurbation) was subjected to lab-scale batch experiments under various conditions of oxygenation and a gradient of methanol addition. The results show that during nitrification, N₂O emissions are positively related to oxygenation (R ² = 0.99). However, compared to the rates of ammonium oxidation, the percentage of emitted N₂O is greater when oxygenation is low (0.5–1 mgO₂ L⁻¹), representing up to 1% of the oxidized ammonium (0.4% on average). During denitrification, the N₂O emission reaches a significant peak when the quantity of methanol allows denitrification of between 66% and 88%. When methanol concentrations lead to a denitrification of close to 100%, the flows of N₂O are much lower and represent on average 0.2% of the reduced nitrate. By considering these results, we can estimate, the emissions of N₂O during nitrogen removal, at the “Seine Centre” WWTP, to approximately 38 kgN-N₂O day⁻¹.     

Isotopic composition of nitrate-nitrogen as a marker of riparian and benthic denitrification at the scale of the whole Seine River system

Nitrogen budgets established for large river systems reveal that up to 60% of the nitrate exported from agricultural soils is eliminated, either when crossing riparian wetlands areas before even reaching surface waters, or within the rivers themselves through benthic denitrification. The study of nitrogen isotope ratios of riverine nitrates could offer an elegant means to assess the extent of denitrification and thus confirm these budgets, as it is known that denitrification results in a natural ¹⁵N enrichment of residual nitrates. The results reported here, for the Seine river system (France), demonstrate the feasibility of this isotopic approach at the scale of large watersheds. On the basis of in situ observations carried out in a large storage reservoir in the upstream Seine catchment (Der Lake), where intensive benthic denitrification occurs, as well as on the basis of laboratory experiments of denitrification under controlled conditions, it is shown that the isotopic discrimination associated with benthic denitrification is minimal ( of NO₃-N ranging from –1.5 to –3.6), probably because the rate-limiting step of the process consists of nitrate diffusion through the water-sediment interface. Riparian denitrification on the contrary, when it implies nitrate reduction during convective transfer through reducing environements, causes a much more significant isotopic enrichment of ¹⁵N of residual nitrate ( about –18). The authors report measurements of nitrogen isotopic composition of nitrate from rivers of various stream orders in the Seine river system under summer conditions. Anomalies in the data with respect to the values expected from the mixture of the various sources of nitrate are here attributed to riparian denitrification. However, the authors show that because of the patchy distribution of actively denitrifying riparian zones within the drainage network, the isotopic signature conferred to residual nitrate in river water intrinsically provides only a minimum estimate of the extent of denitrification.     

Nitrogen Fixation by the Photosynthetic Sulfur Bacterium Chlorobium phaeobacteroides from Lake Kinneret

N₂ fixation by Chlorobium phaeobacteroides from Lake Kinneret was dependent on ammonia concentration and light intensity. In the thermocline of Lake Kinneret, N₂ fixation and photosynthesis were low. It was concluded that the bacteria do not contribute significantly to the organic nitrogen load of the lake.     

Denitrification and total N losses from an irrigated sandy-clay loam under maize–wheat cropping system

Denitrification and total N losses were quantified from an irrigated field cropped to maize and wheat, each receiving urea at 100 kg N ha^-1. During the maize growing season (60 days), the denitrification loss measured directly by acetylene inhibition-soil cover method amounted 2.72 kg N ha^-1 whereas total N loss measured by ¹⁵N balance was 39 kg ha^-1. Most (87%) of the denitrification loss under maize occurred during the first two irrigation cycles. During the wheat growing season (150 days), the denitrification loss directly measured by acetylene inhibition-soil cover and acetylene inhibition-soil core methods was 1.14 and 3.39 kg N ha^-1, respectively in contrast to 33 kg N ha^-1 loss measured by ¹⁵N balance. Most (70-88%) of the denitrification loss under wheat occurred during the first three irrigation cycles. Soil moisture and NO ₃ ^- -N were the major factors limiting denitrification under both crops. Higher N losses measured by ¹⁵N balance than C₂H₂ inhibition method were perhaps due to underestimation of denitrification by C₂H₂ inhibition method and losses other than denitrification, most probably NH₃ volatilization.     

HOT WATER EXTRACTABLE PHOSPHORUS—AN INDICATOR OF NUTRITIONAL STATUS OF PERIDINIUM CINCTUM (DINOPHYCEAE) FROM LAKE KINNERET (ISRAEL)?1

The intracellular levels of hot water extractable and total phosphorus were determined in the dinoflagellate Peridinium cinctum. f. westii (Lemm.) Lef. for natural samples from the bloom in Lake Kinneret and from laboratory cultures. Amounts of phosphorus (P) in the hot water fraction, relative to total cellular phosphorus, were similar in lake Peridinium and in cells grown in high ambient orthophosphate (P_i) media (3–6 mg P · l^?1). The absolute amounts of hot water extractable P in natural cell and those cultured at lower P_i concentrations (0.02–0.05 mg P · 1^?1) were similar, although average P_i in lake water were 4 μg · l^?1. Under most growth conditions the hot water extract contained approximately equal amounts of molybdate reactive phosphorus (MRP) and non-MRP. Short chain (6–9 units) polyphosphates (mol wt 630–950) probably constituted the bulk of the non-MRP pool, which was hydrolysable by alkaline phosphatase and may serve as a precursor for a more permanent P store. Intracellular P levels and distribution were not directly dependent on external P_i concentrations but may be determined by the N:P atomic ratio or overall external ionic milieu. Peridinium grown in low ambient P_i released significant amounts of non-MRP compounds. In Lake Kinneret, for at least most of the bloom period, Peridinium does not appear to be limited by P supply.     

In situ and experimental evidence of the influence of turbulence on cell division processes of Peridinium cinctum forma westii (Lemm.) Lefèvre

Comparison of the diel variations of the wind intensity and the division rate (DR) of Peridinium cinctum forma westii (Lemm.) Lefèvre in situ reveals that the intensity of the wind blowing throughout the whole day does not affect the DR. On the other hand, a strong inhibitory effect is noticed when the wind episodes occur during the time period 18.00–02.00 h. Systematic hourly sampling and staining of Peridinium cells showed that nuclear division takes place between 23.00 and 02.00 h and is completed before cytokinesis begins. Thus the time period 18.00–02.00 h corresponds to the premitotic and mitotic phases of cell division, and the turbulence generated by the wind affects the process of nuclear division.

The relationship between water turbulence and the DR of Peridinium which was observed in Lake Kinneret (Israel) has been checked under experimental conditions. Peridinium was grown without shaking, with continuous rotary shaking and with intermittent shaking at 100 r min^-1. The specific growth rate (k), generation time (G) and mortality rate were followed and compared. The results obtained confirm the facts observed in situ and clarify some aspects. Intermittent shaking of 2 h day^-1 during the dark phase reveals the inhibitory effect of agitation on nuclear division. Continuous shaking causes a high rate of cell mortality. Shaking during the light period does not affect the division process.

The effect of turbulence on the DR of Peridinium explains why the timing of the bloom in Lake Kinneret is a function of the duration and intensity of the mixing period in the lake.     

Hydrological variability, organic matter supply and denitrification in the Garonne River ecosystem

1. Groundwater nitrate contamination has become a worldwide problem as increasing amounts of nitrogen fertilisers are used in agriculture. Alluvial groundwater is uniquely juxtaposed between soils and streams. Hydrological connections among these subsystems regulate nutrient cycling. 2. We measured denitrification using an in situ acetylene‐block assay in a nitrate‐contaminated portion of the Garonne River catchment along a gradient of surface water–ground water mixing during high (snowmelt) and low flow. 3. During high flow (mid‐April to early June) the water table rose an average of 35 cm and river water penetrated the subsurface to a great extent in monitoring wells. Denitrification rates averaged 5.40 μgN₂O L^?1 min^?1 during the high flow period, nearly double the average rate (2.91 μgN₂O L^?1 min^?1) measured during base flow. This was driven by a strong increase in denitrification in groundwater under native riparian vegetation. Nitrate concentrations were significantly lower during high flow compared with base flow. Riparian patches had higher dissolved organic carbon concentrations that were more aromatic compared with the gravel bar patch closest to the river. 4. Multiple linear regression showed that the rate of denitrification was best predicted by the concentration of low molecular weight organic acids. These molecules are probably derived from decomposition of soil organic matter and are an important energy source for anaerobic respiratory processes like denitrification. The second best predictor was per cent surface water, reflecting higher denitrification rates during spring when hydrological connection between surface water and ground water was greatest. 5. Our results indicate that, while denitrification rates in Garonne River alluvium were spatially and temporally variable, denitrification was a significant NO₃ sink during transport from the NO₃‐contaminated floodplain to the river. DOC availability and river–floodplain connectivity were important factors influencing observed spatial and temporal patterns.     

Phytoplankton nutrient deficiency in lakes of the McMurdo dry valleys, Antarctica

1. The influence of inorganic nitrogen and phosphorus enrichment on phytoplankton photosynthesis was investigated in Lakes Bonney (east and west lobes), Hoare, Fryxell and Vanda, which lie in the ablation valleys adjacent to McMurdo Sound, Antarctica. Bioassay experiments were conducted during the austral summer on phytoplankton populations just beneath the permanent ice cover in all lakes and on populations forming deep-chlorophyll maxima in the east and west lobes of Lake Bonney. 2. Phytoplankton photosynthesis in surface and mid-depth (13 m) samples from both lobes of Lake Bonney were stimulated significantly (P < 0.01) by phosphorus enrichment (2 μM) with further stimulation by simultaneous phosphorus plus NH₄⁺ (20 μM) enrichment. Similar trends were observed in deeper waters (18 m) from the east lobe of Lake Bonney, although they were not statistically significant at P < 0.05. Photosynthesis in this lake was never enhanced by the addition of 20 μM NH₄⁺ alone. Simultaneous addition of phosphorus plus nitrogen stimulated photosynthesis significantly (P < 0.01) in both Lake Hoare and Lake Fryxell. No nutrient response occurred in Lake Vanda, where activity in nutrient-enriched samples was below unamended controls; results from Lake Vanda are suspect owing to excessively long sample storage in the field resulting from logistic constraints. 3. Ambient dissolved inorganic nitrogen (DIN) (NH4⁺+ NO₂^?+ NO₃^?): soluble reactive phosphorus (SRP) ratios partially support results from bioassay experiments indicating strong phosphorus deficiency in Lake Bonney and nitrogen deficiency in Lakes Hoare and Fryxell. DIN : SRP ratios also imply phosphorus deficiency in Lake Vanda, although not as strong as in Lake Bonney. Particulate carbon (PC): particulate nitrogen (PN) ratios all exceed published ratios for balanced phytoplankton growth, indicative of nitrogen deficiency. 4. Vertical nutrient profiles in concert with low advective flux, indicate that new (sensu Dugdale & Goering, 1967) phytoplankton production in these lakes is supported by upward diffusion of nutrients from deep nutrient pools. This contention was tested by computing upward DIN : SRP flux ratios across horizontal planes located immediately beneath each chlorophyll maximum and about 2 m beneath the ice (to examine flux to the phytoplankton immediately below the ice cover). These flux ratios further corroborated nutrient bioassay results and bulk DIN : SRP ratios indicating phosphorus deficiency in Lakes Bonney and Vanda and potential nitrogen deficiency in Lakes Hoare and Fryxell. 5. Neither biochemical reactions nor physical processes appear to be responsible for differences in nutrient deficiency among the study lakes. The differences may instead be related to conditions which existed before or during the evolution of the lakes.     

Nitrification, denitrification, and nitrate ammonification in sediments of two coastal lagoons in Southern France

Summary Seasonal and diurnal variations in sediment-water fluxes of O₂, NO ₃ ^– , and NH ₄ ⁺ as well as rates of nitrification, denitrification, and nitrate ammonification were determined in two different coastal lagoons of southern France: The seagrass (Zostera noltii) dominated tidal Bassin d'Arcachon and the dystrophic Etang du Prévost. Overall, denitrification rates in both Bassin d'Arcachon (<0.4 mmol m^–2 d^–1) and Etang du Prévost (<1 mmol m^–2 d^–1) were low. This was mainly caused by a combination of low NO ₃ ^– concentrations in the water column and a low nitrification activity within the sediment. In both Bassin d'Arcachon and Etang du Prévost, rates of nitrate ammonification were quantitatively as important as denitrification.Denitrification played a minor role as a nitrogen sink in both systems. In the tidal influenced Bassin d'Arcachon, Z. noltii was quantitatively more important than denitrification as a nitrogen sink due to the high assimilation rates of the plants. Throughout the year, Z. noltii stabilized the mudflats of the bay by its well- developed root matrix and controlled the nitrogen cycle due to its high uptake rates. In contrast, the lack of rooted macrophytes, and dominance of floating macroalgae, made nitrogen cycling in Etang du Prévost more unstable and unpredictable. Inhibition of nitrification and denitrification during the dystrophic crisis in the summer time increased the inorganic nitrogen flux from the sediment to the water column and thus increased the degree of benthic-pelagic coupling within this bay. During winter, however, benthic microalgae colonizing the sediment surface changed the sediment in the lagoon from being a nitrogen source to the over lying water to being a sink due to their high assimilation rates. It is likely, however, that this assimilated nitrogen is liberated to the water column at the onset of summer thereby fueling the extensive growth of the floating macroalgae, Ulva sp. The combination of a high nitrogen coupling between sediment and water column, little water exchange and low denitrification rates resulted in an unstable system with fast growing algal species such as phytoplankton and floating algae.     

Measurement of denitrification in rivers: an integrated, whole reach approach

Rivers are believed to play an important role in nitrogen removal via denitrification. Unfortunately, there are few data quantifying these processes in situ, primarily due to methodological constraints. We have developed a new approach for estimating denitrification in rivers at the whole reach scale and have applied this approach to three small rivers, the Millstone River in central New Jersey, and the Iroquois River and Sugar Creek in northwest Indiana–northeast Illinois (USA). The approach is based on measuring the change in dissolved N₂ concentration as a parcel of water moves downstream. Two volatile, non-reactive tracers (propane and isobutane) were co-injected, and the rate of change in the ratio of these gases was used to calculate a first-order transfer rate of N₂ (K_N2) to correct for loss of the gas to the atmosphere. Nitrogen removal via denitrification ranged between 0.27 ± 1.21 mmol N m^–2 h^–1 in Sugar Creek during May 2000 and 15.81 ± 2.51 mmol N m^–2 h^–1 in the Millstone River during March 2001. This approach could permit testing of factors that are believed to control denitrification at the reach scale, such as nitrate concentration, discharge, temperature, and water residence time, and could provide a clearer picture of nitrogen transformations in rivers.     

Net nitrogen mineralization,nitrification and CO2 production in alternating moisture conditions in an unfertilized low-humus sandy soil from the Sahel

In this study the rates of net mineralization, net immobilization and net nitrification have been quantified under laboratory conditions in a sandy low-humus soil from a semi-arid region, in absence of plant growth. Incubation experiments were carried out under constant humidity and under alternating wet and dry conditions to simulate field conditions during the rainy season. The ammonium and nitrate content of the incubates were determined and their CO₂ production measured.The rate of net mineralization at field capacity was 0.6 kg N ha^–1d^–1 during the first 40 days and decreased to 0.06 kg N ha^–1d^–1 after 400 days. This rate was twice as high on wet days under alternating wet and dry conditions. The rate of net nitrification during alternating wet and dry conditions was also higher (1.9 kg N ha^–1d^–1) than at constant field capacity (1.3 kg N ha^–1d^–1) until the ammonium was almost completely depleted. These rates of net mineralization and net nitrification are in agreement with field observations.Net immobilization did not occur in the experiments, unless glucose was added to the soil.The data on CO₂ production and net mineralization showed that the C/N ratio of the degraded material was around 9 or below. It is much lower than the ratio of total carbon over total nitrogen in the soil. This indicates that microorganisms and compounds high in nitrogen were mineralized. Certainly after about 30 days the only growth taking place is based on turnover of material of the microbial biomass itself.A decrease in the amount of inorganic nitrogen was observed upon drying of the soil, while it returned to the original content after rewetting. It is postulated that this might be due to temporary uptake of nitrogen in an inorganic form in microorganisms as part of their osmoregulation.     

Effects of growth conditions of acetate utilization byRhodopseudomonas palustris isolated from a freshwater lake

Rhodopseudomonas palustris, a purple non-sulfur bacterium, was recently found throughout the water column in Lake Kinneret. It was demonstrated to be of a versatile nature, growing under both aerobic and anaerobic conditions at different light intensities. A comparison of C-acetate uptake byR. palustris andChlorobium phaeobacterioides, a green sulfur bacterium, showed that, under identical growth conditions, C-acetate assimilation byR. palustris was greater. Furthermore, C-acetate uptake forR. palustris was greater than C−CO₂ uptake at all light intensities. Depending on the prevailing conditions, acetate can be used byR. palustris as both an electron donor and carbon source. Malate synthase was used as an indicator of activity of the glyoxylic acid cycle. It was found that enzyme activity was higher (i.e., acetate was used mainly as a carbon source) under anaerobic conditions, in the dark, or in the absence of HCO ₃ ⁻ . Acetate was used preferably as an electron donor under photosynthetic microaerophillic conditions.     

Seasonal Rotifer Dynamics in the Long-term (1969–2002) Record from Lake Kinneret (Israel)

Long-term (1969–2002) data record of biomass distribution of rotifers in Lake Kinneret is combined with previously published information on their metabolic activity and newly calculated population dynamics parameters to synthesize a model of their seasonal dynamics in Lake Kinneret. Nineteen rotifer species were recorded in routine samples collected in Lake Kinneret (Israel) in 7 offshore (deeper than 5 m), stations, at 12 discrete depths during 1969–2002. Organisms were sorted and counted (including external egg carrying females), biomass was measured and calculated for the entire lake stock (g_w.w m⁻²; mg l⁻¹). Rates of grazing, respiration and production were measured experimentally at three different temperature ranges. Results were extrapolated to the lake community for months with similar temperatures. Rotifera comprised 7% of total zooplankton biomass in Lake Kinneret whilst Cladocera and Copepoda 58 and 35% respectively. Rotifers were found to be more abundant during December–June and decline in summer months. Monthly (1969–2001) means indicated total grazing capacity of rotifers as 11%, respiration as 9% and production as 3.7% of the total zooplankton metabolic activity. Positive relations were indicated between rotifer and small bodied cladoceran numerical concentrations. Population growth models suggest that rotifers are not food limited in Lake Kinneret but that fish predation plays an important role in regulating abundance in spring-summer and fall.     

Nutrient loading and drainage channel response in a ricefield system

The distribution of photosynthetic bacteria in Lake Kinneret and their contribution to primary productivity was investigated. Both Chlorobium phaeobacteroides and Rhodopseudomonas palustris were detected from mid-June through October in 1987 and reached a maximum of 7 × 10⁶ cells/ml in the metalimnion by mid-August. Chlorobium density was greater towards the end of the bloom period, when the thermocline was exposed to higher sulfide concentrations and lower light intensities. Rhodopseudomonas peaked earlier in the season, when less sulphide was present and light irradiation was higher. This was the first time that R. palustris was observed in Lake Kinneret; in addition to its presence at the thermocline, it was found also at the epilimnion and hypolimmon. The contribution of photosynthetic bacteria to the total primary productivity of the lake was small (∼1%). Low but perceptible rates of N₂ fixation in situ were also demonstrated by photosynthetic bacteria present in the metalimnion.     

Field Measurements of Photosynthesis of Umbilicarious Lichens in Winter

The photosynthetic performance of Lasallia pustulata and Umbilicaria spodochroa was measured both in the field on granitic rock in southern Norway and in the laboratory under controlled conditions of light and temperature. In the field thallus temperatures varied between ? 2 and + 5 °C during the daylight period in January 1994. In situ water contents were between 50 and 400% d.wt. in L. pustulata and between 100 and 500% d.wt. in U. spodochroa. The lichens were active during the whole period of investigation. Photosynthetic rates reached 13.03 μmol CO₂ g Chl^?1 s^?1 in L. pustulata and 5.56 μmol CO₂ g Chl^?1 s^?1 in U. spodochroa. Ice formation on the thallus surface did not impede CO₂ exchange. Light was mainly the limiting factor, as could be confirmed by the laboratory experiments. In general, habitat conditions never provided optimum photosynthetic rates but photosynthetic carbon balance was positive during 4 of the 5 days investigated. The coldest day was photosynthetically almost as efficient as the warmest day during this period.