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     

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.     

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     

Distribution of dissolved organic carbon and dissolved fulvic acid in mesotrophic Lake Biwa, Japan

The dissolved organic carbon (DOC) concentrations in mesotrophic Lake Biwa were determined by a total organic carbon (TOC) analyzer, and DOC molecular size distributions were determined by size exclusion chromatography (SEC) using a fluorescence detector at excitation/emission (Ex/Em) levels of 300/425 nm with the eluent at pH 9.7. The fluorescence wavelengths for detection were chosen from the result of excitation–emission matrix spectrometry (EEM) analysis for dissolved fulvic acid (DFA) extracted from Ado River (peak A, Ex/Em = 260–270/430–440 nm; peak B, Ex/Em = 300–310/420–430 nm). Ado River DFA was eluted with a retention time (RT) of 7.4–8.9 min and the apparent molecular weight was estimated at 22–87 kDa based on the elution curve for the spherical protein molecular weight standard. A DFA peak eluted at the same retention time as Ado River DFA also appeared in all the samples of Lake Biwa water. From the linear relationship between the peak areas with an RT of 7.4–8.9 min by SEC analysis and DOC values of DFA by TOC analysis of a series of DFA samples (r² = 0.9995), the concentrations of DFA in the lake water were roughly calculated. DFA was distributed within the range 0.25–0.43 mg C l⁻¹ and accounted for 15%–41% of DOC, with the highest ratios observed at a depth of 70 m in August and the lowest at 2.5 m in May.     

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.     

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     

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.     

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     

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.     

Three-dimensional fluorescence as a tool for investigating the dynamics of dissolved organic matter in the Lake Biwa watershed

Quantitative and qualitative characterizations of dissolved organic matter (DOM) were carried out at the watershed level in central Japan by measuring dissolved organic carbon (DOC) concentration and the three-dimensional excitation–emission matrix (3-D EEM). DOC concentration was low (mean 37 ± 19 µM C) in the upstream waters, whereas, in general, it increased toward the downstream areas (mean 92 ± 47 µM C). Significant variations in DOC concentration were detected among rivers and channels. DOC concentration in the epilimnion of Lake Biwa increased during the summer period and decreased during the winter period. The lake hypolimnion has lower DOC concentration (mean 87 ± 7 µM C) compared with the epilimnion (107 ± 15 µM C). Fulvic acid (FA)-like substances in the DOM were directly characterized by 3-D EEM. The fluorescence peak for upstream DOM was found in regions with longer wavelengths (excitation/emission 386 ± 6/476 ± 5 nm) compared with downstream and lake DOM (351 ± 12/446 ± 15 nm and 341 ± 6/434 ± 6 nm, respectively). The DOC concentration is correlated with fluorescence peak intensity of FA-like substances in DOM in river waters. Such a relationship was not found in lake DOM. A blueshift of the fluorescence peak from upstream to lake DOM was observed. A decrease in fluorescence intensities was also detected during the summer period. These results may suggest that the degradation of FA-like substances in DOM occurs from natural solar irradiation. Protein-like fluorescence was significantly detected in the lake epilimnion during the summer period. A linear relationship between DOC concentration and protein-like fluorescence indicated that an autochthonous input of DOM gave rise to the increase in DOC concentration in the lake epilimnion during the summer. These results may suggest that the 3-D EEM can be used as a tool for the investigation of DOM dynamics at the watershed level with concurrent measurement of DOC concentration and the fluorescence properties of fulvic acid-like and protein-like substances.     

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     

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.     

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.     

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.     

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     

Annual cycle of circulations in Lake Biwa, part 1: model validation

We perform a series of model studies of the physical environment in Lake Biwa. In part 1, provided here, we formulate and validate a numerical model which consists of three-dimensional momentum and continuity equations for a Boussinesq fluid under the hydrostatic approximation in a rotating frame, the advective–diffusive equation for water temperature, and the full nonlinear equation of state for freshwater. Water motion is driven by heat flux and wind forcing which are evaluated from the time series of meteorological parameters observed at weather stations around and on the lake using bulk formulae. This model for the first time accurately simulates the annual and interannual variations found in observed water temperature and velocity fields in the 2004–2006 hindcast experiment. A current system of cyclonic (counterclockwise) and anticyclonic (clockwise) circular gyres appears above the seasonal thermocline during a stratified season, and a bottom flow appears along the eastern slope to sink cold dense water from the shallow eastern region to the deep lake bottom during an unstratified season. Full overturning of the water column during winter causes bottom water temperature to change interannually depending on the severity of winter. These successful results endorse the benefits of the model when investigating the physical environment of the lake over longer timescales.     

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.     

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.     

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.     

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.