Effect of temperature and soluble reactive phosphorus on abundance of Aphanizomenon flos-aquae (Cyanophyceae)

Filament density of Aphanizomenon flos‐aquae (Lemmerm.) Ralfs, water temperature and soluble reactive phosphorus (SRP) were measured from April to August in 1993–1996 in Lake Barato, Hokkaido, Japan. In addition, growth characteristics and internal phosphorus (P) utilization of Aph. flos‐aquae were evaluated under P limitation at three temperatures (15, 20 and 25?C) to clarify the role of internal accumulated P for its growth in the incubation experiment. The filament density was highest in early July 1994, when SRP concentration had not yet decreased and the water temperature was high. These are important factors favoring an increase in abundance of this species in L. Barato. During batch culture, the time course of the stationary phase was shortest at 25?C and longest at 15?C; the cellular C:P molar ratio was 111 under P sufficiency and increased eight‐ to 12‐fold under P limitation. As the C:P ratio was significantly higher in the decreasing phase at 15?C, Aph. flos‐aquae may be more adaptable to Plimitation at 15?C than at 20?C and 25?C. However, the low temperatures did not favor the abundance of Aph. flos‐aquae in 1996. This indicates that the filament density of Aph. flos‐aquae decreases before it reaches the maximum value for some reason under P limitation in L. Barato.     

Importance of interspecific competition in the abundance of Aphanizomenon flos-aquae (Cyanophyceae)

The population dynamics of Aphanizomenon flos-aquae var. klebahnii Elenk. (Cyanophyceae) was studied in an artificial pond for 32 months from May 2002 to December 2004. Our previous in vitro study showed that the lower limits of water temperature and pH for its growth are within the ranges of 11°–14°C and 7.1–7.4, respectively, and it appeared that these findings are applicable to the emergence and disappearance of Ap. flos-aquae in the pond. Based on the change in the water temperature, the emergence of Ap. flos-aquae in 2004 was expected to be in late April, whereas emergence occurred after a 1-month lag period. During this period, Ankistrodesmus falcatus (Corda) Ralfs (Chlorophyceae) dominated the phytoplankton assemblage, which raised the possibility that the growth of Ap. flos-aquae was restricted by the existence of An. falcatus. We conducted mixed cultures of Ap. flos-aquae and An. falcatus at four temperatures (14°, 19°, 24°, and 29°C). After 18 days of incubation, An. falcatus dominated at 14° and 29°C whereas Ap. flos-aquae dominated at 19°C. This result indicates that a slightly higher water temperature than the growth threshold value is needed for Ap. flosaquae to outcompete An. falcatus, which agrees with the field observation. Contrary to the results of the mixed culture, the summer phytoplankton assemblage was dominated by Ap. flos-aquae, and the population of An. falcatus was less or almost absent. This variation seemed to be partly caused by the difference in nutrient conditions; concentrations of dissolved inorganic nitrogen and phosphorus in the pond were far lower than those in the culture medium. The lack of nitrogen fixation of An. falcatus seemed to be a growth disadvantage during the summer when the concentration of dissolved inorganic nitrogen was low.     

The formation and degradation of cyanobacterium Aphanizomenon flos-aquae blooms: the importance of pH, water temperature, and day length

Investigation of annual changes in phytoplankton community structure in a small artificial eutrophic pond was carried out from May 2002 to April 2003. A heavy bloom of Aphanizomenon flos-aquae var. klebahnii Elenk. (Cyanobacteria) persisted in most of the water column from June to the end of October. In November, the A. flos-aquae bloom suddenly crashed and green algae were predominant until the end of spring. Weekly monitoring suggested strong involvement of the changes in abiotic factors in the cyanobacterial bloom degradation. To clarify the effects of pH, water temperature, and day length on the growth of A. flos-aquae, laboratory batch experiments were conducted. The results showed that A. flos-aquae could not grow below pH 7.1 and 11°C, and the growth tended to be suppressed under a 10L:14D photoperiod. pH, water temperature, and day length are vital factors in the growth of A. flos-aquae and, additionally, grazing by cyclopoid copepods also seemed important in bloom collapse.     

Harvesting of Aphanizomenon flos-aquae Ralfs ex Born. & Flah. var. flos-aquae (Cyanobacteria) from Klamath Lake for human dietary use

In western cultures, certain cyanobacteria have beenan accepted source of microalgal biomass for food forabout 30 years, in particular Spirulina(Arthrospira) platensis and S. maxima. Beginning in the early 1980s, another species, Aphanizomenon flos-aquae was adopted for similaruses. This is harvested from Upper Klamath Lake, thelargest freshwater lake system in Oregon. In 1998 theannual commercial production of Aphanizomenonflos-aquae was about 1 × 10⁶ kg. Since thisspecies is not cultured like Spirulina inoutdoor ponds or raceways, it requires very differentprocedures for harvesting and processing. These arereviewed here and include extensive off-lake screensor on-lake barges, which dewater and concentrate thecells. Other procedures, such as those for removal ofdetritus and mineral materials, and those formonitoring and reducing the amounts of certaincontaminant cyanobacteria, which can producecyanotoxins, have also become important in qualitycontrol and marketing.     

Notes on the growth and sporulation of a natural population of Aphanizomenon flos-aquae

Aphanizomenon flos-aquae contributed regularly to the summer phytoplankton in Kinnego Bay, a highly eutrophic part of Lough Neagh. Growth of the alga was variable in different years, and was apparently related to the duration of depletion of dissolved nitrate-nitrogen in the water. Although four growth pulses occurred during the study period, sporulation was only observed during one of these, when the population was very dense although evenly distributed down the water column. It seems that Aphanizomenon overwintered in Kinnego Bay as vegetative filaments and that production of akinetes was not necessary for perennation of the species. The observed sporulation in 1973 may have been induced by the high pH values caused by dense phytoplankton crops; no other measured environmental factors were particularly unfavourable to growth of Aphanizomenon at this time.     

Release of phosphorus from sediments in Lake Biwa

Two sulfur-mediated reactions are resulting in the eutrophication of Lake Biwa, Japan. The iron (II) phosphate mineral vivianite is dissolving in sulfide-enriched sediments that in places results in porewater concentrations of phosphate exceeding 3 mg l⁻¹. The dissolution of phosphate is evident in profiles of total phosphorus where zones of dissolution and a zone of precipitation in the most oxic surface sediments are visible. At times sulfate reduction in these surface sediments results in pH values as high as 9.9, which can dissolve phosphate adsorbed to iron (III). This release of phosphorus from sediments is at least partially responsible for the recent appearance of blue-green algal blooms. Received: August 4, 2000 / Accepted: March 19, 2001     

Sources of dissolved methane in Lake Biwa

The spatial distribution and seasonal variation in the concentration and carbon isotopic composition of dissolved methane in a river–lake ecosystem were studied in Lake Biwa, Japan, and its tributary rivers. Methane concentrations in all subsystems examined were supersaturated with respect to the atmosphere. The epilimnion showed higher concentrations of dissolved methane than the hypolimnion in the pelagic zone. Peak methane concentrations were observed at the thermocline. The largest amount of methane in the pelagic water column was recorded at the end of a stagnant period, at which the bottom water of the sublittoral zone (30m in depth) exhibited increased methane concentration. Transect observation of dissolved methane revealed three methane peaks at different water depths in the lake, and river water and the sediments in littoral and sublittoral zones were suggested to be the corresponding sources. Water at the river mouth was replete with dissolved oxygen but also contained a high concentration of methane. The present results suggest that river water and littoral sediment are potential sources of dissolved methane in Lake Biwa, and other sources, such as internal waves, are responsible for increased methane in the pelagic zone at the end of stagnant periods. Carbon stable isotope analysis indicated that there were different sources of dissolved methane, although it was difficult to identify the origins due to high variation of the isotopic composition of methane from different sources.     

Spatial changes of phytoplanktonic size spectra in Lake Biwa

The degree of eutrophication in Lake Biwa was examined from the characteristics of spatial distribution in size spectra computed by a size spectra analyser with an optical fiber sensor. Two types of size spectra could be recognised: (1) size distribution only presenting a peak in a size less than about 20 m in equivalent diameter and (2) size distribution with two peaks in size spectra.The second pattern with two peaks was observed in the size spectra from the south basin, the sites near the central part and the southern part of the north basin, and seemed to be associated with advanced eutrophic conditions. In addition, the degree of eutrophication along the east side of the south basin was examined from changes in size spectra and from hydrological conditions. We inferred that the eutrophic condition around the central sites of the north basin is influenced by the water current from the gyre.     

Effects of zooplankton on the sinking flux of organic carbon in Lake Biwa

To clarify the roles of zooplankton in the sedimentation of seston from the epilimnion, the sinking flux of particulate carbon was measured along with primary production rate and zooplankton biomass from July 1996 to October 1997 at a pelagic site in the north basin of Lake Biwa. During the study period, the flux varied seasonally from 66 to 510 mg C m⁻² day⁻¹ and was low in summer when zooplankton, composed mainly of Eodiaptomus japonicus and Daphnia galeata, were abundant. Simple correlation analysis revealed that the sinking flux correlated neither with the primary production rate nor with the amount of sestonic carbon above the sediment trap. However, the particle elimination rate, estimated as the difference between the primary production rate and the sinking flux, correlated positively with the zooplankton biomass. These results suggest that zooplankton play a substantial role in decreasing the sinking flux in Lake Biwa. Received: March 6, 2000 / Accepted: October 7, 2000     

Urea degradation by picophytoplankton in the euphotic zone of Lake Biwa

The ability of photoautotrophic picoplankton Synechococcus to degrade urea was examined in the euphotic zone of Lake Biwa. Samples were divided into pico (0.2–2.0 μm) and larger (>2.0 μm) size fractions by filtration. The rates of urea degradation (the sum of the rates of incorporation of carbon into phytoplankton cells and of liberation of CO₂ into water) measured by radiocarbon urea were 8 and 17 μmol urea m⁻³ day⁻¹ in June and July, respectively, for the picophytoplankton in the surface water, and 196 and 96 μmol urea m⁻³ day⁻¹, respectively for the larger phytoplankton. The rates decreased with depth, somewhat similar to the vertical profiles of the photosynthetic rate. The urea degradation rates were obviously high under light conditions. In daylight, urea was degraded into two phases, carbon incorporation and CO₂ liberation, whereas in the dark it was degraded only into the CO₂ liberation phase. The contribution of picophytoplankton to total phytoplankton in urea degradation was high in the subsurface to lower euphotic layer. Urea degradation activity was higher in the picophytoplankton fraction than in the larger phytoplankton fraction. Shorter residence times of urea were obtained in the upper euphotic zone. The contribution of picophytoplankton to urea cycling was 4% to 35%. The present results suggest that the picophytoplankton Synechococcus is able to degrade urea and effectively makes use of regenerated urea as a nitrogen source in the euphotic layer, and that picophytoplankton play an important role in the biogeochemical nitrogen cycle in Lake Biwa. Received: June 25, 1998 / Accepted: February 10, 1999     

Rotifer community structure in the south basin of Lake Biwa

Seasonal changes in abundance and species composition of rotifers were surveyed at five locations under different physical and chemical conditions in the south basin of Lake Biwa during July 1987–June 1988. Total density of rotifers showed similar seasonal fluctuation with three peaks, although the maximum density showed north (low)-south (high) gradient. Polyarthra spp. (P. vulgaris and P. dolichoptera with low density) dominated except during July–October 1987 and April–May 1988. In the former period the species of Brachionus, Trichocerca, Filinia and Hexarthra, and in the latter, those of Synchaeta, Keratella and Kellicottia had somewhat different proportions in the communities of north and south stations, respectively. The difference in composition during July–October suggests a difference of trophic state between the northern and southern areas in the south basin of Lake Biwa. However, the dominance of Polyarthra and the difference in the composition during April–May 1988 could not be explained by such a difference in trophic state. No critical difference was observed in the community structure of the eastern area of the south basin, where the seasonal fluctuations in nutrient levels and phytoplankton community structure were different from other areas in the south basin. The present results, therefore, suggest that physical and chemical conditions were not effective in controlling the rotifer community structure in the south basin of Lake Biwa.     

On the Occurrence of Aphanizomenon flos-aquae var. Flos-aquae in Fish Ponds

Summer blooms of Aphanizomenon flos-aquae var. flos-aquae accompanied by dense populations of Daphnia were common in fish ponds of the district Blatná in 1950s-1960s. Changes in management (increased stocking densities, decreased usage of superphosphate as fertilizer) made this type of plankton less frequent. The present study documents this trend and suggests explanations.     

Spatial distribution and seasonal changes of pesticides in Lake Biwa,Japan

The spatial distribution and seasonal variation in the concentrations in Lake Biwa of pesticides used in paddy fields were studied. Lake Biwa is the largest lake in Japan and is a recognized water resource for 14 million people in the Kinki district. Samples were collected nine times from April to December 2001 at ten sites within the lake and at the mouths of six influent rivers. Weekly sampling was also carried out at a single site on an effluent river. Among the 20 pesticides analyzed, the detection frequencies in surface water were almost 100% for simetryn, bromobutide, and isoprothiolane; around 75% for molinate and pyroquilon; around 30% for three herbicides and one fungicide; and almost zero for the remaining substances. The maximum concentrations of pesticides detected frequently in the lake were in the range 0.1–0.4µgl^–1. The occurrence of a few pesticides below the thermocline may be explained by thermal stratification and vertical circulation. Although the thermocline suppressed vertical diffusion in spring and summer during pesticide application periods, a few pesticides remaining at the surface of the lake in winter were transported to the hypolimnion by vertical circulation and remained there even after the reestablishment of the thermocline. The half-lives of pesticides in the lake were estimated to be more than a year for simetryn, half a year for bromobutide, 1.5 months for molinate, and 1 month for dimepiperate. The main cause of elimination for molinate and dimepiperate was estimated to be degradation, that for simetryn was outflow, and for bromobutide both degradation and outflow were significant.     

The budget of dissolved trace metals in Lake Biwa,Japan

A comprehensive study on the dynamics of dissolved elements (Mg, Al, Si, P, Ca, V, Cr, Mn, Fe, Ni, Zn, As, Sr, Y, W, and U) in Lake Biwa was carried out using a clean technique. Lake water samples (n = 523) were collected from six stations in the North Basin and three stations in the South Basin. River water samples (n = 178) were collected from 14 major rivers flowing into the North Basin. Rainwater samples (n = 89) were collected at Otsu. The river water was enriched with Mn, Al, Fe, P, and Zn and the rainwater was enriched with Zn, Al, Fe, and Mn compared to North Basin water during winter mixing. The residence times of dissolved species were estimated on the basis of input through the rivers and rain. The residence times for Ca, Mg, and Sr were about 8 years, the same as that for water. Mn, Al, Fe, and Zn showed the shortest residence times (0.05–0.19 year). A budget calculation suggested that more than 60% of the input of dissolved Si, P, V, Cr, Mn, Fe, Ni, and Zn was scavenged and retained in the lake sediments and/or discharged as suspended particles.     

Horizontal distribution of carbon and nitrogen and their isotopic compositions in the surface sediment of Lake Biwa

The horizontal distribution of the abundance and isotopic composition of carbon and nitrogen was studied on surface sediment samples (0–15 cm) collected from the entire area of Lake Biwa, the largest freshwater lake in Japan. As water depth increased, a marked increase in organic matter content was observed at the sampling sites, especially in the western North Basin, characterized by a steep slope. In the northwestern North Basin, which has no major inflowing streams, the sediments contained large amounts of organic matter, suggesting the possibility of lateral transportation of sedimented matter from other places by lake currents. The total amounts of carbon and nitrogen in the top-2 cm of sediment of the entire area of Lake Biwa were estimated to be 9.2 × 10⁴ tC and 1.0 × 10⁴ tN. The δ¹⁵N values in the littoral sediment were low and close to those in the inflowing river sediment, suggesting selective sedimentation of allochthonous organic matter onto the littoral area. In the North Basin, vertical profiles of organic matter content and δ¹³C values of the sediments in the littoral area showed a smaller downward decrease than in the profundal area, whereas δ¹⁵N values decreased with sediment depth in both areas. It was suggested that the littoral sediments contained abundant amounts of allochthonous and relatively refractory organic matter. Further, it was suggested that the autochthonous organic matter originated from primary production deposited mainly on the profundal zone and was easily decomposed in early diagenesis after sedimentation. Received: July 30, 1999 / Accepted: December 10, 1999     

Seasonal variation of dissolved aluminum concentration in harmonic-type Lake Biwa, Japan

Monthly observations performed on a typical harmonic-type lake, Lake Biwa (latitude 35°15′ N, longitude 136°05′ E, Japan), showed that the particulate aluminum concentration varied around a mean value of 0.8 μM, with occasional extraordinarily high values, whereas the dissolved aluminum concentration varied, in the surface layer of the whole lake, between a minimum of 0.01 μM and a maximum of 0.30 μM, depending on the season. Although the variation in dissolved aluminum paralleled the variation in the pH of the lake water, the variation in dissolved aluminum lagged behind the variation in pH by approximately 1 month. A series of laboratory incubation experiments suggested that the supply of dissolved aluminum from, and its removal by, the suspended particulate matter involved a slow, pH-dependent reaction. The stoichiometry and the apparent equilibrium constant of this reaction were evaluated by adopting a zeolite-like structure for the surfaces of the suspended particulates. Received: December 28, 2000 / Accepted: August 22, 2001     

Paddy herbicide inputs in the entire river inflow reaching Lake Biwa, Japan

This study estimated the inputs of four paddy herbicides in the entire river inflow reaching Lake Biwa, the largest lake in Japan, which serves as a water resource for 14 million people. The Uso River and the Hino River, the main contaminated rivers among the inflow rivers, were selected as daily and hourly monitoring sites to provide data on the seasonal trends in the concentration and load of herbicides and to determine the effect of rainfall events on load. The monitoring was also performed four times in 15 inflow rivers. The total input to the lake was calculated from the loads during fine weather conditions and additional loads during rainfall events. The former based on the lumped load from the two rivers and by prorating for the 15 rivers, and the latter was estimated from the relation between precipitation and increased load rate. The annual losses of herbicide from the basin to Lake Biwa were estimated to be 14.5% for bromobutide, 3.0% for pretilachlor, 5.2% for molinate, and 8.8% for simetryn. The loads caused by rainfall events accounted for 9%–18% of the total annual loads.     

Geochemical behavior of trace elements in Lake Biwa

A monthly survey of dissolved concentrations of various trace elements was performed in Lake Biwa. Particulate concentrations of the elements were also measured in early autumn and winter. Based on these results, the geochemical behaviors of trace elements are discussed. The redox-sensitive elements Mn and Fe showed characteristic vertical distribution profiles. Profiles of Mn changed drastically with the progression of the stagnation period. The dynamics of Ba were affected by the redox cycle of Mn. Dissolved V concentration showed a clear seasonal variation. In contrast, dissolved concentrations of Sr, Mo, Cu, Zn, and Ni were almost uniform, i.e., not dependent on the season or the depth. The distribution ratios of these elements between lake water and Mn nodules formed in the lake were calculated to assess their geochemical behaviors.     

Distribution of CH4 in the north basin of Lake Biwa and tributary rivers

To determine the origin of CH₄, the vertical distribution of CH₄ around the thermocline in the north basin of Lake Biwa and the horizontal distribution in the Yasu and Ado rivers were measured. In 1995, CH₄ concentrations (ranging from 200 to 1000 nM) exceeding the saturation level were observed just above or in the thermocline in the pelagic region off the Yasu River. These values were higher than those just below the thermocline and in the hypolimnion. CH₄ concentrations in the mouths of the Yasu and Ado rivers were much higher than at other stations, around 2500 and 2000 nM, respectively. Due to the drought in 1994 when there was no water from the Yasu River flowing into the north basin, CH₄ concentration just above or in the thermocline in the pelagic region off the Yasu River ranged from 49 to 74 nM. It is thus concluded that the high level of CH₄ observed in the mouth of the Yasu River is one of the sources of the high CH₄ concentrations in the pelagic region off the Yasu River. On the other hand, the CH₄ concentration in the pelagic region off the Ado River was about 50 nM. It is thought that water flowing in from the Ado River diffused readily into the lake water because the depth of the lake in the region off the Ado River declines steeply. The maximum concentration of CH₄ in the river mouths indicates that these areas are important sites for clarifying the mechanism of the decrease in dissolved oxygen in Lake Biwa and the Yodo River watershed. Received: October 25, 2000 / Accepted: May 8, 2001     

Effects of temperature, and availability of nitrogen and phosphorus on the abundance of Anabaena and Microcystis in Lake Biwa, Japan: an experimental approach

Under optimal nutrient conditions, both Microcystis sp. and Anabaena sp. isolated from Lake Biwa grew optimally at 28–32°C but differed in maximal growth rates, phosphate uptake kinetics, maximal phosphorus quotas, and growth responses to nitrogen and phosphorus limitation. The maximal growth rates of Microcystis and Anabaena were 1.6 and 1.25 divisions day⁻¹, respectively. With phosphate and nitrate in the growth-limiting range, the growth of Microcystis was optimal at an N : P ratio of 100 : 1 (by weight) and declined at lower (nitrogen limitation) and higher (phosphorus limitation) ratios. In contrast, Anabaena growth rates did not change at N : P ratios from 1000 : 1 to 10 : 1. Starting with cells containing the maximal phosphorus quota, Microcystis growth in minus-phosphorus medium ceased in 7–9 days, compared with 12–13 days for Anabaena. The phosphate turnover time in cultures starved to their minimum cell quotas was 7.9 min for Microcystis and 0.6 min for Anabaena. Microcystis had a higher K _s (0.12 μg P l⁻¹ 10⁻⁶ cells) and lower V _max (9.63 μg P l⁻¹ h⁻¹ 10⁻⁶ cells), than Anabaena (K _s 0.02 μg P l⁻¹ h⁻¹ 10⁻⁶ cells; V _max 46.25 63 μg P l⁻¹ h⁻¹ 10⁻⁶ cells), suggesting that Microcystis would not be able to grow well in phosphorus-limited waters. We conclude that in spite of the higher growth rate under ideal conditions, Microcystis does not usually bloom in the North Basin because of low availability of phosphorus and nitrogen. Although Anabaena has an efficient phosphorus-uptake system, its main strategy for growth in low-phosphorus environments may depend on storage of phosphorus during periods of abundant phosphorus supply, which are rare in the North Basin. Received: July 31, 2000 / Accepted: October 18, 2000