Geochemistry of Lake Biwa sediments revisited

Extensive chemical data for the Lake Biwa sediments were examined with statistical factor analysis to find any correlation patterns among elements and among samples. Based on those patterns, a series of x-y correlation plots of the concentrations of selected elements was constructed to see in detail how the concentrations of elements vary among different sample groups and explain the observed variation patterns.     

Long-term light environment variability in Lake Biwa and Lake Kasumigaura, Japan: modeling approach

Light environment variability was investigated in the two Japanese lakes Biwa and Kasumigaura, which offer a broad range of optical conditions in the water bodies due to their diverse morphometries and limnological characteristics. To elucidate their light environments, Secchi depths (SDs) were related to long-term monitored datasets of concentrations of optically active substances (OASs) using two approaches based on statistical and mechanistic models. A good estimate for the nonphytoplanktonic suspended solids (NPSS) concentration gained using a monthly factor ?? (which represents the phytoplanktonic portion in total suspended solids) from a long-term analysis was utilized to develop robust models. Using the mechanistic model, the contribution of each OAS to the SD can be understood and investigated in more detail than possible with a statistical approach, but the statistical model yields better results in terms of SD prediction. Based on the results of an analysis of the contribution of each OAS to the SD, it was clear that NPSS was the component that exerted the most influence on the light environments in the two lakes; in this respect, this study agrees with other studies that show the importance of suspended inorganic particles as the main contributor to the SD in inland waters. Using ANOVA, we analyzed how specific inherent optical properties may have changed spatially and temporally, and the results indicated that the temporal (monthly) effect was primarily responsible for the loss of accuracy in the models. In addition, the ANOVA analysis suggested that grouping the data improved the predictive performances of the statistical models. Finally, we concluded that combining the two models yields the most reliable results in terms of SD prediction and determining the contribution of each OAS to the SD at present.     

Asian great lakes, especially Lake Biwa

Synopsis The geological and biological history of Asian great lakes, especially Lake Biwa in Japan, are reviewed. The origins and affiliations of endemic flora and fauna are interpreted in light of current understanding of Lake Biwa. Recent historical changes in the lake, including detrimental impacts on native fauna and water quality are summarized. The social and cultural values associated with Asian lakes are reviewed, and contrasted to those of lakes elsewhere. In general Asian great lakes are smaller than those elsewhere, with the notable exception of Lake Baikal. Furthermore, Asian lakes are typically more eutrophic, with higher primary productivity. Societal values in China and other Asian countries have tended to favour nutrient enrichment and increased cultured fish production from lakes. Asian societies, however, have long attached important artistic, poetic and cultural values to their lakes.Paper from the Canadian Society of Zoologists symposium Great Lakes of the World, organized by David L.G. Noakes.     

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.     

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     

Genetic identification of larvae and juveniles reveals the difference in the spawning site among Cyprininae fish species/subspecies in Lake Biwa

To understand the differences in the spawning sites among Cyprininae fishes in Lake Biwa, we conducted periodic sampling of larvae and juveniles at three sites (irrigation ditch, St. 1; river, St. 2; and satellite lake, St. 3). On the basis of species/subspecies identification by using RAPD analysis, we examined the species composition of the larvae and juveniles at these three sampling sites. The number of specimens was 616, 68, and 117 at St. 1, St. 2, and St. 3, respectively. Based on morphological and genetic identification, the specimens were found to include nine fish species/subspecies, namely, Carassius auratus grandoculis, Carassius cuvieri, Carassius auratus langsdorfii, Cyprinus carpio, Sarcocheilichthys sp., Silurus asotus, Oryzias latipes, Odontobutis obscura obscura, and Rhinogobius sp. The species composition at the three sites also differed. Among the Cyprininae fishes, C. auratus grandoculis, C. auratus langsdorfii, and Cyprinus carpio were found in abundance at St. 1; C. cuvieri was not collected from St. 1 but was found at the other two sites, particularly St. 3. Among the other fishes, Rhinogobius sp. was collected at St. 1 and St. 3, whereas the other four occurred only at St. 1. These results suggest that the selection of spawning sites by C. cuvieri differs to a certain extent from that of the other Cyprininae fishes, and the irrigation ditch in the lake is an important habitat for the larvae and juveniles of native fish species.     

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.     

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.     

Effect of temperature on growth of the cyanobacterium Aphanizomenon flos-aquae in Lake Biwa and Lake Yogo

The water bloom‐forming cyanobacterium Aphanizomenon flos‐aquae Ralfs ex Bornet et Flahault (Nos‐tocales, Cyanophyceae) appeared in Lake Biwa and Lake Yogo in 1999 for the first time. The morphological characteristics were described using natural samples. In contrast to the other water bloom‐forming cyanobacteria such as Microcystis and Anabaena in Lake Biwa and Lake Yogo, the small summer population of A. flos‐aquae is apt to grow in winter, suggesting the low temperature preference or tolerance of this species. In order to clarify the effect of temperature on the growth, culture experiments were conducted using an axenic strain isolated from Lake Biwa. The strain could grow at above 8°C with an optimum temperature ranging from 23 to 29°C, and survived even at 5°C for at least 25days under low light conditions. Although these results confirmed the ability of the bloom formation during late autumn and winter, it is still unclear why the Aphanizomenon bloom occurred at temperatures of ca 10°C in December and not immediately after the disappearance of Microcystis and/or Anabaena bloom during autumn.     

Biotic and abiotic methane releases from Lake Biwa sediment slurry

To determine the rate and mechanism of CH₄ production in Lake Biwa sediment, slurry was prepared and incubated. Surface sediment (sed) slurry (1.5–6cm) showed a CH₄ release rate (4.9–9.5nmolg-dry-sed^–1 day^–1) higher than that observed in the 5- to 10-cm sediment slurry (0.2–2nmolg-dry-sed^–1 day^–1). Methane release from the surface (1.5–6cm) sediment slurry was biotic and was inhibited by addition of 2-bromoethanesulfonate (BES, an inhibitor of CH₄ production), whereas that from 5- to 10-cm sediment slurry was abiotic. The addition of BES, HNO₃, and O₂ showed no effect on the CH₄ release rate from the 5- to 10-cm sediment slurry. In addition, tracers (NaH¹³CO₃, ¹³CH₃COONa) were not incorporated into the released CH₄. However, ¹³C of CH₄ released from the 5- to 10-cm sediment slurry (–74.0 ± 0.6) indicated that this CH₄ was produced by bacterial metabolism in the past, stored by adsorption on the surface of clay minerals in the sediment, and then released abiotically by desorption from the sediment slurry as a result of a decrease in hydraulic pressure and CH₄ concentration in the pore water. This CH₄ stored by adsorption could be extracted by autoclaving. In the sediment below 5cm, bacterial activity for CH₄ production ceased, possibly because of the limitated availability of H₂. To clarify the mechanism of CH₄ production in the sediment, biotic CH₄ production and the abiotic CH₄ release found here should be estimated separately.     

First observation of incomplete vertical circulation in Lake Biwa

Limnology - Vertical circulation reaches the bottom of the deepest (> 90 m) part of the northern basin of Lake Biwa, a warm monomictic lake, during winter (January to March)...     

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.     

Annual cycle of circulations in Lake Biwa, part 2: mechanisms

We investigate the mechanisms of circulations in Lake Biwa using model results that are driven by the momentum and heat fluxes evaluated from meteorological parameters observed around and on the lake. The kinetic energy budget shows that the wind stress over the lake, which has positive (negative) curl over the northern (southern) region of the lake throughout the year, generates the current system of multiple gyres in the stratified season (May–January), while the thermal forcing makes a minor contribution. This mechanism is quite different from the previous picture, in which the thermal forcing was dominant in forming the gyre system during the heating season. In the cooling season, the overturning of the lake occurs not in a simple one-dimensional (vertical) process, but in a time-dependent three-dimensional one. Cold water that is formed in the shallow eastern region descends the bottom slope to the deep lake in December to March while such a descent hardly occurs in the western region. A large shallow area (small heat capacity) as well as intense cooling in the eastern region effectively produces cold water. Wind again plays an important role by intensifying the cooling through its higher speed and by driving slope current. Further, the fluctuating wind direction causes significantly time-dependent behaviors of the slope current or bottom water formation.     

Seasonal succession of zooplankton in the north basin of Lake Biwa

To examine the seasonal succession of the entire zooplankton community in Lake Biwa, zooplankton biomass (on an areal basis) and its distribution patterns among crustaceans, rotifers and ciliates were studied in the north basin from April 1997 to June 1998. Seasonal changes in phytoplankton and population dynamics of Daphnia galeata were also examined to assess food condition and predation pressure by fish. From March to November, crustaceans dominated zooplankton biomass, but rotifers and ciliates were dominant from December to February. Among crustaceans, Eodiaptomus japonicus was the most abundant species, followed by D. galeata. Zooplankton biomass increased from January to a peak in early April, just before the spring bloom of phytoplankton, then decreased in mid-April when mortality rate of D. galeata increased. From mid-June, zooplankton increased and maintained a high level until the beginning of November. During this period, both birth and mortality rates of D. galeata were relatively high and a number of rotifer and crustacean species were observed. However, their abundances were very limited except for E. japonicus which likely preys on ciliates and rotifers. In Lake Biwa, food sources other than phytoplankton, such as resuspended organic matter from the sediments, seems to play a crucial role in zooplankton succession from winter to early spring, while zooplankton community seems to be regulated mainly by fish predation from summer to fall.     

Seasonal characteristics of surface water fulvic acids from Lake Biwa and Lake Tankai in Japan

Seasonal characteristics of surface water fulvic acids (FAs) isolated from Japanese clear-water lakes were investigated. Qualitative changes in Lake Biwa and Lake Tankai FAs were determined and compared. Although the relative molecular weights determined by high-performance size-exclusion chromatography did not change remarkably, the elemental compositions, E₆₀₀^1% E_{600}^{1% } values and ¹H- and ¹³C-NMR spectral properties varied with the season. Both the H/C and N/C ratios for Lake Biwa FAs tended to be higher than those for Lake Tankai FAs, but O/C ratios were lower and decreased from spring to winter. The E₆₀₀^1% E_{600}^{1% } values suggested that Lake Biwa FAs contained lower levels of unsaturated structures than Lake Tankai FAs, and the amounts of these structures increased in winter. The ¹H and ¹³C NMR spectra indicated that Lake Biwa FAs are richer in saturated aliphatic chains, especially in spring. Overall, Lake Biwa FAs exhibited clearer seasonal changes in these characteristics than Lake Tankai FAs, suggesting that the seasonal variation may depend on changes in aquatic microbial activities.     

The production-to-respiration ratio and its implication in Lake Biwa,Japan

Production-to-respiration (P:R) ratio was estimated at an offshore site of Lake Biwa in order to examine whether the plankton and benthic community is subsidized with allochthonous organic carbon, and to clarify the role of this lake as potential source or sink of carbon dioxide. The respiration rate of protozoan and metazoan plankton was calculated from their biomass and empirical equations of oxygen consumption rates, and that of bacterioplankton was derived from their production rate and growth efficiency. In addition, the carbon mineralization rate in the lake sediments was estimated from the accumulation rate of organic carbon, which was determined using a ²¹⁰Pb dating technique. On an annual basis, the sum of respiration rates of heterotrophic plankton was comparable to net primary production rate measured by the ¹³C method. However, when the mineralization rate in the lake sediments was included, the areal P:R ratio was 0.89, suggesting that Lake Biwa is net heterotrophic at the offshore site with the community being subsidized with allochthonous organic carbon. Such a view was supported by the surface water pCO₂ that was on average higher than that of the atmosphere. However, the estimate of net CO₂ release rate was close to that of carbon burial rate in the sediments. The result suggests that the role of Lake Biwa in relation to atmospheric carbon is almost null at the offshore site, although the community is supported partially by organic carbon released from the surrounding areas.     

Effects of seston on ultraviolet attenuation in Lake Biwa

We examined the attenuation of underwater ultraviolet (UV) radiation and photosynthetically available radiation (PAR) in Lake Biwa, Japan, at offshore and inshore sites and under contrasting stratification and mixing regimes. There were large spatial differences in the water column transparency to both wavebands, despite little change in concentrations of dissolved organic carbon (DOC). The 1% of surface irradiance depth varied from 0.3 to 2.7 m at 305 nm, from 0.8 to 6.3 m at 380 nm, and from 2.3 to 12.8 m for PAR. Both PAR and UV transparency declined abruptly in the South Basin of the lake when a typhoon caused the resuspension of sediments. The water column ratio of UV to PAR increased by 30% at all stations over the course of a 3-week sampling period associated with the general increase in phytoplankton concentrations. At several sites, the diffuse attenuation coefficient for UV radiation deviated substantially from that predicted from UV-DOC models. A significantly positive linear relationship was found between UV attenuation (K _d determined with a profiling UV radiometer) and the beam attenuation coefficient at 660 nm as measured by transmissometer. These results indicate that scattering and absorption by particulate matter can reduce UV transparency to below that inferred from DOC concentrations, and that current UV-exposure models should be modified to incorporate this effect. Received: March 21, 2001 / Accepted: August 17, 2001     

Size-dependent phytoplankton responses to atmospheric forcing in Lake Biwa

Size-dependent changes in chlorophyll a and uptake of inorganiccarbon (C) and nitrogen (N) by phytoplankton were measured inthe shallow South Basin of Lake Biwa, before and during a periodof typhoons (high winds). The latter period was characterizedby complete mixing of the water column, a major decline in underwaterirradiance, and a transient increase in dissolved reactive Nand phosphorus (P). Nutrient concentrations, seston N:P ratiosand uptake rates indicate that P and not N limited phytoplanktonover the whole study period. The typhoon-induced increase inphosphorus supply resulted in Blackman-type (increased C- andN-specific growth rates) and Liebig-type (increased biomassyield) responses by the phytoplankton. Picoplankton were dominantin the relatively stable and clear water column prior to thetyphoons, but were rapidly outgrown by larger cells during andafter wind-induced mixing. The slower response of picoplanktonindicates lower maximum intrinsic growth rates and photosyntheticefficiency. These observations are contrary to the view thatpicoplankton have higher light-harvesting abilities and growthrates than larger cells. Typhoon-induced mixing stimulated theshift to fast-growing, dim light-adapted larger cells. The resultsquestion the allometric paradigm of increasing growth ratesand light-capturing efficiency with decreasing cell size, butare consistent with the oceanographic view that large-celledphytoplankton control the major fluctuations in biomass andprimary productivity, while picoplankton account for a comparativelystable background productivity.     

Dynamics of methane in mesotrophic Lake Biwa,Japan

As a part of a core project of IGBP (International Geosphere-Biosphere Programme), distribution, production, oxidation and transport processes of methane in bottom sediments and lake water in a mesotrophic lake (Lake Biwa) have been studied with special reference to the spatial heterogeneity of each process. In this study, we attempted to synthesize previously reported results with newly obtained ones to depict the methane dynamics in the entire lake. The pelagic water column exhibited subsurface maxima of dissolved methane during a stratified period. Transect observation at the littoral zone suggested that horizontal transportation may be a reason for the high methane concentration in epilimnion and thermocline at the offshore area. Tributary rivers and littoral sediments were suggested to be the source. Observations also showed that the internal wave caused resuspension of the bottom sediment and release of methane from the sediment into the lake water. The impact of the internal waves was pronounced in the late stage of a stratified period. The littoral sediment showed much higher methanogenic activity than the profundal sediments, and the bottom water of the littoral sediments had little methanotrophic activity. In the profundal sediment, most of the methane that diffused up from the deeper part was oxidized when it passed through the oxic layer. Active methane oxidation was also observed in the hypolimnetic water, while the lake water in the epilimnion and thermocline showed very low methane oxidation, probably due to the inhibitory effect of light. These results mean a longer residence time for methane in the epilimnion than in the hypolimnion. Horizontal inflow of dissolved methane from the river and/or littoral sediment, together with the longer residence time in the surface water, may cause the subsurface maxima, which have also been observed in other lakes and in the ocean.     

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