Algal nitrogen fixation in the tropics

Summary a)Nitrogen fixation in rice fields. Nitrogen-fixing blue-green algae grow abundantly in tropical regions and are particularly common in paddy fields. Their possible role in the nitrogen accumulation of soil has been studied. The most vigorous nitrogen-fixing blue-green algae have been assessed for use as green manure in rice fields and favorable effects have been reported in India and other countries. b)Nitrogen fixation by algae in water. The planktonic blue-green algae occur abundantly at certain time of the year in sea water and lake water, and some of them are known to be nitrogen fixers. Certain Japanese species of blue-green algae can withstand high temperatures including ten nitrogen-fixing species from hot-spring waters. c)Nitrogen fixation by symbiotic blue-green algae. Certain species of blue-green algae form associations with other organisms such as fungi, liverworts, ferns and seed plants. The relationship between these two organisms is on one occasion commensal and on others symbiotic. Certain symbiotic blue-green algae are provided with the ability to fix the atmospheric nitrogen.     

The effect of nutrient enrichment on nitrogen fixation activity in the Bay of Quinte,Lake Ontario

Nitrogen fixation was measured in situ by the ¹⁵N tracer technique in the Bay of Quinte, Lake Ontario, and three lake enclosures with different nutrient enrichment. The fixation rates in the Bay were low but detectable during the summer season. The fixation activities were found to be correlated with the presence of nitrogen-fixing blue-green algae and the distribution of the algal species in the water was affected by nitrate enrichment. The study showed that, with the addition of nitrate, species not able to fix atmospheric nitrogen became predominant. However, in the absence of external nitrogen (i.e., nitrate), species able to fix nitrogen became dominant. Phosphorus enrichment alone did result in higher N-fixation rates in the water and, without the addition of phosphorus, the fixation rates are lower and fluctuate throughout the season, presumably dependent on the availability of phosphorus in the water.A comparison between the ¹⁵N-isotopic method and the acetylene reduction method is reported and the factors involved in the variations between these two methods are discussed.     

A numerical model of nitrogen flxation and its application to Lake Valencia,Venezuela*

SUMMARY. 1. A numerical model for calculation of daily and annual nitrogen fixation in lakes is presented. The model is based on empirically-derived equations for the rates of nitrogen fixation by heterocysts (nitrogen-fixing cells) in relation to light and on functions for the vertical and tetnporal distributions of heterocysts and light in a lake. 2. Applications of the model to Lake Valencia, Venezuela, between December 1980 and December 1981 indicated that nitrogen fixation is largely a surface phenomenon in this lake: 80% of diurnal fixation occurred within 1m of the water surface. 3. Nitrogen fixation is largely restricted to periods of lake stratification, when the phytoplankton have sufficient light for growth, but dissolved inorganic nitrogen is scarce. Nitrogen fixation was maximal late in the stratification period of 1981: 85 % of fixation occurred within the last 3 months of the 9-month period. 4. The annual nitrogen fixation in Lake Valencia is 26 kg ha^?1, which is comparable to the nitrogen fixation in temperate eutrophic lakes with seasonal blue-green algal blooms. However, nitrogen fixation accounted for only 23% of the total nitrogen supply to Lake Valencia in 1981.     

Algal nitrogen fixation on the lava field of Heimaey,Iceland

Summary Nitrogen fixation (C₂H₂ reduction) by blue-green algae occurring on the juvenile lava field of Heimaey, Iceland was examined both in the laboratory (potential at 20° C and 39° C) and in the field, three and a half years after the volcanic eruption.Already at this early stage of colonization representatives of unicellular and filamentous heterocystous and non-heterocystous blue-green algae were commonly observed. The predominating algae were Nostoc sp. (20° C) and Schizothrix sp. — Microcoleus chthonoplastes, (39° C), the former often in association with the protonemata-rhizoids of moss plants.The potential for nitrogen fixation was recorded at an average rate of 109.2 (20° C) and 138.1 (39° C) ng N g^-1 h^-1 in soil collected from localities randomly distributed over the lava field.Tests for nitrogen fixation performed in situ revealed significant fixation activities in all the eleven localities subject to examination. The activities ranged from 2.8 to 63.4 (mean 21.5) ng N g^-1 h^-1 and 1.9 to 17.7 (mean 7.9) ng N cm^-2 h^-1.All the nitrogen fixation data noted imply that blue-green algae contribute a substantial part of the nitrogen input to the lava. Further, it was found that material incubated under micro-aerophilic conditions exhibited considerably enhanced nitrogenase activity.The role of nitrogen-fixing blue-green algae in general and Nostoc muscorum in particular in being suitable as pioneering organisms preparing the bare lava for ingress of other plants is also discussed.     

The Utricularia-Cyanophyta association and its nitrogen-fixing capacity

Utricularia inflexa Forsk. growing in a shallow lake near Dar es Salaam, Tanzania was found to be abundantly associated with blue-green algae and other microorganisms, though blue-green algae were not found in the lake water. The epiphytes occurred both on the outer surfaces of the macrophyte and inside its traps. Seven genera of the Cyanophyta were observed, amongst which Anabaena was the most abundantly and consistently associated.Both epiphytes on the outer surfaces of the macrophyte and those inside mature traps fixed nitrogen in situ. Mature parts of the macrophyte fixed nitrogen at higher rates than immature parts, indicating the former was associated with a greater abundance of nitrogen fixers. The Utricularia-Cyanophyta association showed a nitrogen fixation (acetylene reduction) rate of 2012 nmol C₂H₄ g^–1 dry weight h^–1 or, in terms of lake area, of 4 500 nmol C₂H₄ m^–2 h^–1. The relative contributions of blue-green algae and bacteria were not determined.It is proposed that the association may involve a degree of physiological interdependence and that the association has potential as a biofertilizer for rice.Based on an M.Sc. thesis by G. M. Wagner submitted to the Department of Botany, University of Dar es Salaam, 1983.Based on an M.Sc. thesis by G. M. Wagner submitted to the Department of Botany, University of Dar es Salaam, 1983.     

低温对氯化钠胁迫下蓝藻固氮活性的影响

低温加剧氯化钠对蓝藻固氮的抑制,营养液中氯化钠浓度增高时,抑制程度更甚.能源受限(暗处理和加抑制剂时的光合受抑,N_2和Ar的厌氧下呼吸代谢受阻)和氧下固氮酶受到伤害时,低温处理使氯化钠对蓝藻固氮的抑制进一步加剧.在能源和还原剂供应,合成固氨酶蛋白的物质基础(如CO_2和N_2的加合).光合作用正常进行的条件得到改善和保证,以及供应CO_2、外源蔗糖和氮氧加合时,低温加剧氯化钠对蓝藻固氮的抑制程度明显变小.     

Phytoplankton ecology of the Lake of Menteith,Scotland

The results discussed in this paper represent the first seasonal ecological study carried out on the phytoplankton of the Lake of Menteith. All measured nutrients reached maximum levels during the winter, with silicate showing particularly high concentrations (up to 85 µg at Si l^–1). During the summer period phosphate, nitrate and silicate showed almost complete exhaustion in surface waters. The lake water was consistently alkaline, never falling below pH 7, while the alkalinity ranged from 20 to 24 mg CaCO₃ l^–1. Generally, the nutrient status of the main inflow had a rapid effect on the water quality of the lake.The region of the lake under investigation showed no thermal stratification at any period of the year, although continuous thermal gradients were recorded in the winter. The continual circulation of the water mass probably prevented oxygen saturation from falling below 77% even following a large phytoplankton bloom and subsequent decomposition.From an examination of net phytoplankton samples the Lake of Menteith could be described as blue-green or blue-green/diatom in nature. From the quantitative study, large pulses of Melosira, Asterionella and Fragilaria were recorded in the spring. The disappearance of the species appears to be related to silicate limitation. The summer growth of Asterionella may have been promoted by a nitrogen source other than nitrate and nitrite, both of which were reduced to critical levels. This alternative source of combined nitrogen may have been contributed by nitrogen-fixing algae in the lake. Three species of Anabaena were recorded, all of which produced large populations during the year.Department of Botany, The University of GlasgowPresent Address: Department of Biology, College of Science, University of Sulaimaniyah, Sulaimaniyah, Iraq     

The Effects of Moisture on Acetylene Reduction by Mats of Blue-green Algae in Sub-tropical Grassland

During the dry winter months desiccation is the main factorpreventing nitrogen fixation by mats of blue-green algae inkikuyu lawns. In the wetter summer months a few days of sunshinewithout rain will dry out the algal mats and depress nitrogenfixation unless the algae are protected by swards of grass.Dry mats resume nitrogen fixation within an hour of becomingdamp and attain their normal rate of fixation within 5 h. Maximumfixation rates occur at 100 per cent relative humidity and withsoil moisture contents of 22 to 42 per cent. Dry algal matsare unable to take up sufficient water as vapour (even in saturatingconditions) to permit resumption of acetylene reduction. Algal-dominatedsoils show greater water retention than adjacent bare soils.     

Effect of Rice Plants on Nitrogenase Activity of Flooded Soils

In samples of flooded soil containing blue-green algae (cyanobacteria), the presence of rice plants did not influence the nitrogenase activity of the algae. Nitrogenase activity of heterotrophic bacteria was enhanced by the presence of rice plants, but this activity was not affected by changes in plant density. The rate of nitrogen fixation in the rhizosphere, however, varied significantly among the 16 rice varieties tested. A simple method was devised to test the nitrogen-fixing activity in the root zone of rice varieties, and data were obtained showing marked differences in the activities of the 16 varieties. In tests of two varieties with dissimilar rates of nitrogen fixation in their rhizospheres, the variety which had the greater root weight and lesser shoot weight and which supported greater methane formation had the greater nitrogenase activity.     

Decomposition of Blue-Green Algal (Cyanobacterial) Blooms in Lake Mendota, Wisconsin

Decomposition of natural populations of Lake Mendota phytoplankton dominated by blue-green algae (cyanobacteria) was monitored by using oxygen uptake and disappearance of chlorophyll, algal volume (fluorescence microscopy), particulate protein, particulate organic carbon, and photosynthetic ability (¹⁴CO₂ up-take). In some experiments, decomposition of ¹⁴C-labeled axenic cultures of Anabaena sp. was also measured. In addition to decomposition, mineralization of inorganic nitrogen and phosphorus were followed in some experiments. Decomposition could be described as a first-order process, and the rate of decomposition was similar to that found by others using pure cultures of eucaryotic algae. Nitrogen and phosphorus never limited the decomposition process, even when the lake water was severely limited in soluble forms of these nutrients. This suggests that the bacteria responsible for decomposition can obtain all of their key nutrients for growth from the blue-green algal cells. Filtration of lake water through plankton netting that removed up to 90% of the algal biomass usually did not cause a similar decrease in oxygen demand, suggesting that most of the particulate organic matter used for respiration of the decomposing bacteria was in a small-particle fraction. Short-term oxygen demand correlated well with the particulate chlorophyll concentration of the sample, and a relationship was derived that could be used to predict community respiration of the lake from chlorophyll concentration. Kinetic analysis showed that not all analyzed components disappeared at the same rate during the decomposition process. The relative rates of decrease of the measured parameters were as follows: photosynthetic ability > algal volume > particulate chlorophyll > particulate protein. Decomposition of ¹⁴C-labeled Anabaena occurred at similar rates with aerobic epilimnetic water and with anaerobic sediment, but was considerably slower with anaerobic hypolimnetic water. Of the various genera present in the lake, Aphanizomenon and Anabaena were more sensitive to decomposition than was Microcystis. In addition to providing a general picture of the decomposition process, the present work relates to other work on sedimentation to provide a detailed picture of the fate of blue-green algal biomass in a eutrophic lake ecosystem.     

Pseudomonas aeruginosa Chemotaxis Associated with Blooms of N2-Fixing Blue-Green Algae (Cyanobacteria)

Pseudomonas aeruginosa (Schroeter) Migula, a numerically significant bacterium found during N₂-fixing blooms of the blue-green algae (cyanobacteria) Anabaena sp. in the Chowan River, North Carolina, was chemotactically attracted to amino acids when tested in a radioassay. The bacterium was labeled with ³²P_i, and the disintegrations per minute determined by liquid scintillation counting were proportional to the number of cells accumulating in microcapillaries containing amino acids. Positive chemotaxis was observed toward all of the amino acids tested, although the degrees of response varied. Since many nitrogen-fixing blue-green algae secrete nitrogenous compounds, this attraction may be instrumental in establishing a symbiotic relationship between this bacterium and blue-green algae in freshwater.     

The effect of ammonium nitrogen on fixation of elemental nitrogen inAlnus andMyrica

Summary 1. A substantial formation of nodules occurred on plants ofAlnus glutinosa andMyrica gale grown in water culture with different levels of ammonium nitrogen labelled with N¹⁵ present in the culture solution. The nodules tended to be fewer but larger than on plants in solution free of combined nitrogen.2. The nodules continued to fix atmospheric nitrogen despite the presence of ammonium nitrogen in the rooting medium, though fixation per unit weight of nodule tissues formed was somewhat lower than in nitrogen-free solution. Among other possible reasons this could have been due to a substitution of ammonium nitrogen for elemental nitrogen at the nitrogen-fixing centres of the nodule, but evidently this does not occur to any great extent.3. In Alnus but not in Myrica fixation per plant was considerably enhanced in the presence of a low level of ammonium nitrogen, owing to greater nodule development. At higher ammonium levels, in excess of the plants' requirements, fixation per plant was still of comparable order to that in nitrogen-free solution, but now only represented some 24 to 45 per cent of the total nitrogen accumulated by the plants.4. The results suggest that under field conditions some fixation of atmospheric nitrogen will always be associated with nodules present.     

Nitrogenase activity,amino acid pool patterns and amination in blue-green algae

Summary The free amino acid pools in the nitrogen-fixing blue-green algae Anabaena cylindrica, A. flos-aquae and Westiellopsis prolifica contain a variety of amino acids with aspartic acid, glutamic acid and the amide glutamine being present in much higher concentrations than the others. This pattern is characteristic of that found in organisms having glutamine synthetage/glutamate synthetase [glutamine amide-2-oxoglutarate amino transferase (oxido-reductase)] as an important pathway of ammonia incorporation. Under nitrogen-starved conditions the level of acetylene reduction (nitrogen fixation) and the glutamine pool both increase but the free ammonia pool decreases, suggesting that ammonia rather than glutamine regulates nitrogen fixation.Glutamine synthetase has been demonstrated in Anabaena cylindrica using the -glutamyl transferase assay and also using a biosynthetic assay in which P_i release from ATP during glutamine synthesis was measured. The enzyme (-glutamyl transferase assay) is present in nitrogen-fixing cultures and activity is higher in aerobic than in microaerophilic cultures. Ammonium-grown cultures have lowest levels of all and activity in the presence of nitrate-nitrogen (150 mg nitrogen 1^-1) is lower than in aerobic cultures growing on elemental nitrogen. Ammonium-nitrogen and nitrate-nitrogen have no effect on glutamine synthetase in vitro. Glutamate synthetase also operates in nitrogen-fixing cultures of Anabaena cylindrica.     

Phosphorus availability and nitrogenase activity in aquatic blue-green algae

Metabolically active phosphorus-starved cultures of blue-green algae assimilate ³²P rapidly in the light and in the dark. The uptake of phosphorus results in a rapid (within 15 min) stimulation in acetylene reduction by Anabaena cylindrica, A. flosaquae, Anabacnopsis circuiaris and Chlorogloea fritschii, with a response being obtained to less than 5 μg/1 of phosphorus. Uptake of phosphorus also causes a rapid increase in respiration in the dark but not in photo respiration, and the size of the cellular ATP pool and the ¹⁴CO₂ fixation rate both increase more slowly. The metabolism of phosphorus-sufficient cells, which assimilate phosphorus more slowly, shows little response when phosphorus is provided. Excess phosphorus is stored in the vegetative cells of blue-green algae as polyphosphate bodies which may form within 60 min of adding phosphorus to phosphorusstarved cells and which serve as a source of phosphorus for the algae when exogenous phosphorus is limiting. Preliminary results from Scottish waters suggest that urban effluents are important sources of available-phosphorus for algal growth and that the levels entering fresh waters from agricultural land are, per unit volume, lower. In both types of water the levels of available-phosphorus are rather similar to the levels of orthophosphate-phosphorus present. Most detergents tested serve as a source of phosphorus for nitrogen-fixing blue-green algae and cause a rapid stimulation in reduction when added to phosphorus-starved cultures. Of the detergents assayed, the biological types were richest in available phosphorus. The addition of detergents may result in a rapid increase in number of polyphosphate bodies present in the algae. Detergents in general also contain an inhibitor of algal metabolism. Whether a stimu-lation or an inhibition occurs depends on the quantities of detergent added and on whether or not the alga is phosphorus-deficient.     

Nitrogenase Activity in Relation to Intracellular Organisms in Sphagnum Mosses

Electron microscopic studies of Sphagnum lindbergii (Schimp.) and S. riparium (Ångstr.) have revealed the presence of intracellular organisms such as blue-green algae, green algae, bacteria and fungi. Nitrogenase activities of these Sphagnum mosses were found to be related mainly to the presence of intracellular Nostoc filaments. The appearance of nitrogen-fixing blue-green algae within bryophytes is thus not restricted to liverworts. The association is likely to be of ecological importance as it seems to occur in very acid habitats generally lacking blue-green algae. Possible interrelations between the moss, the blue-green algae and different types of bacteria are discussed.     

低温对氯化钠胁迫下蓝藻固氮活性的影响

低温加剧氯化钠对蓝藻固氮的抑制,营养液中氯化钠浓度增高时,抑制程度更甚.能源受限(暗处理和加抑制剂时的光合受抑,N_2和Ar的厌氧下呼吸代谢受阻)和氧下固氮酶受到伤害时,低温处理使氯化钠对蓝藻固氮的抑制进一步加剧.在能源和还原剂供应,合成固氨酶蛋白的物质基础(如CO_2和N_2的加合).光合作用正常进行的条件得到改善和保证,以及供应CO_2、外源蔗糖和氮氧加合时,低温加剧氯化钠对蓝藻固氮的抑制程度明显变小.     

Does N2 fixation meet the nitrogen requirements of heterocystous blue-green algae in shallow eutrophic lakes?

Summary Phytoplankton net carbon uptake and nitrogen fixation were studied in two shallow, eutrophic lakes in South Sweden. Ranges of diurnal net carbon uptake were estimated by subtracting 24-h respiration rates corresponding to 5–20% of P _max, respectively, from daytime carbon uptake values. total nitrogen requirement of the phytoplankton assemblage was determined from the diurnal net carbon uptake, assuming a phytoplankton C:N ratio of 9.5:1. Nitrogen supplied by nitrogen fixation only occasionally corresponded to the demands of the total phytoplankton assemblage. When heterocystous algae made up a substantial proportion (10%) of the total phytoplankton biomass, nitrogen fixation could meet the requirements of heterocystous blue-green algae on c. 50% of the sampling occasions. Nitrogen deficiencies in heterocystous algae were most probably balanced by the simultaneous or sequential assimilation of dissolved inorganic nitrogen. It was concluded that uptake of ammonium or nitrate, regenerated from lake seston and sediment, is the main process by which growth of phytoplankton is maintained during summer in the lake ecosystems studied.     

The role of nitrogen as a growth limiting factor in the eutrophic Lake Vesijärvi,southern Finland

The significance of nitrogen for algal growth was studied in Lake Vesijärvi in 1979 and 1980 by algal bioassay, using Selenastrum capricornutum and Anabaena cylindrica as test organisms. Nitrogen limited the growth of Selenastrum for the major part of the investigation period, while phosphorus seemed to be the most limiting factor for Anabaena. This difference was reflected in the in situ succession of phytoplankton. As the ratio of inorganic nitrogen to phosphate phosphorus became smaller, nitrogen-fixing blue-green algae became dominant. Nitrogen fixation was greatest at the beginning of July, coinciding with maximum heterocyst numbers.     

Nitrogen Turnover in Marine and Brackish Habitats: I. Nitrogen Fixation

Potential nitrogen-fixing genera were found to be abundant intwo natural populations of blue-green algae, one from a rockycoast and the other from a sand-dune slack. ¹⁵N studies confirmedthat these populations fixed nitrogen in the laboratory andin the field. Preliminary quantitative data on Fixation in thefield suggest that the algae contribute appreciable quantitiesof fixed nitrogen to the environments in which they occur.     

Effect of Interactions Among Algae on Nitrogen Fixation by Blue-Green Algae (Cyanobacteria) in Flooded Soils

Nitrogen fixation (C₂H₂ reduction) by algae in flooded soil was limited by interactions within the algal community. Nitrogen fixation by either indigenous algae or Tolypothrix tenuis was reduced severalfold by a dense suspension of the green alga Nephrocytium sp. Similarly, interactions between the nitrogen-fixing alga (cyanobacterium) Aulosira 68 and natural densities of indigenous algae limited nitrogen-fixing activity in one of two soils examined. This was demonstrated by developing a variant of Aulosira 68 that was resistant to the herbicide simetryne at concentrations that prevented development of indigenous algae. More nitrogen was fixed by the resistant variant in flooded soil containing herbicide than was fixed in herbicide-free soil by either the indigenous algae or indigenous algae plus the parent strain of Aulosira. Interference from indigenous algae may hamper the development of nitrogen-fixing algae introduced into rice fields in attempts to increase biological nitrogen fixation.