Long-term effects of successive Ca(OH)2 and CaCO3 treatments on the water quality of two eutrophic hardwater lakes

1. Whole-lake experiments were conducted in two hardwater lakes (Halfmoon and Figure Eight) in Alberta, Canada, to investigate the effectiveness of repeated lime (slaked lime: Ca(OH)₂ and/or calcite: CaCO₃) treatments (5–78 mg L^–1) for up to 7 years.
2. Randomized intervention analysis of intersystem differences between the experimental and three reference lakes demonstrated a decline in euphotic total phosphorus and chlorophyll a concentrations in the experimental lakes after repeated lime treatments.
3. After the second lime application to Halfmoon Lake, mean winter total phosphorus release rates (TPRR) decreased to < 1 mg m^–2 day^–1 compared with 3.6 mg m^–2 day^–1 during the winter after initial treatment. In the final year of lime application, mean summer TPRR decreased to 4.5 mg m^–2 day^–1 compared with 7.6 mg m^–2 day^–1 in the pre-treatment year.
4. Mean macrophyte biomass declined and species composition was altered at 1 and 2 m depths in Figure Eight Lake during lime application. Over the first 6 years of treatment, macrophyte biomass at 2 m declined by 95% compared with concentrations recorded during the initial treatment year. In the last year of the study, macrophyte biomass at 2 m reached initial treatment concentrations, which coincided with the greatest water transparency. Over the treatment period, macrophyte species shifted from floating to rooted plants.
5. Multiple lime applications can improve water quality in eutrophic hardwater lakes for periods of up to 7 years.     

Lime treatment and its effects on the chemistry and biota of hardwater eutrophic lakes

1. The main focus of this study was to investigate the effects of single and multiple moderate doses of lime (slaked lime, Ca(OH)₂, and/or calcite, CaCO₃) on eutrophic hardwater lakes. This information would contribute to strategies to manage phytoplankton and macrophyte biomass in eutrophic lakes.
2. Water chemistry and biota were monitored for up to 7 years after initial lime treatment and results were compared with reference systems.
3. Complementary studies investigated the effect of lime on macrophytes in ponds, irrigation canals and microcosm experiments.
4. When water pH was kept within its natural range (≤ 10), single and multiple lime applications to lakes and ponds controlled macrophyte biomass, without negatively affecting invertebrate communities.
5. Single lime treatments at moderate dosages of lakes and ponds resulted in variable and mostly temporary changes in chlorophyll a (chl a ) and phosphorus (P) concentration. Although sediment P release was reduced in single-dose lakes during the first winter following treatment, reductions appeared temporary.
6. Multiple treatments of lakes and ponds were effective at reducing both chl a and P concentrations over longer periods. Mean winter P release rate was also reduced after initial treatment.
7. In laboratory studies, sediment cores were incubated with eight different treatments to assess P release. Redox-sensitive treatments were no more effective at lowering total P concentration in overlying water than some redox-insensitive treatments. Lime reduced total P concentrations, but was not as effective as treatments with alum.
8. The use of lime in managing macrophyte and phytoplankton biomass in shallow, hardwater lakes and ponds may be preferable over other treatments, because lime is economical and non-toxic as long as pH is kept within a natural range.     

Response of plankton communities to whole-lake Ca(OH)2 and CaCO3 additions in eutrophic hardwater lakes

1. The impact of whole-lake lime (slaked lime, Ca(OH)₂, and/or calcite, CaCO₃) addition on plankton communities was evaluated in eutrophic hardwater lakes on the North American Boreal Plain.
2. Two lakes received a single treatment of lime (Ca(OH)₂ at 74 or 107 mg L^–1), two lakes received multiple treatments with Ca(OH)₂ and/or CaCO₃ (5–78 mg L^–1), and four lakes were untreated and served as reference systems.
3. Over the long-term (> 1 year), phytoplankton biomass was reduced in multiple-dose lakes, but not in single-dose lakes. Cyanobacteria typically dominated the algal community in the years before, during and after lime treatment in both single- and multiple-dose lakes.
4. In the single-dose lakes, randomized intervention analysis showed no significant change in the biomass of zooplankton after lime addition.     

Phosphorus release from sediments in hardwater eutrophic lakes: the effects of redox-sensitive and -insensitive chemical treatments

1. Phosphorus (P) release from bottom sediments is an important source of nutrient enrichment in many lakes in sedimentary basins, such as those in western Canada. On the Boreal Plain, sediment P release is particularly strong during periods of seasonal anoxia.
2. In this study, the effects of reduction–oxidation (redox)-sensitive and -insensitive chemicals on P release were examined in sediment cores collected from three eutrophic lakes.
3. Contrary to expectations, redox-sensitive treatments were no more effective at lowering total phosphorus (TP) in sediment cores than some redox-insensitive treatments. Redox-sensitive treatments with FeCl₃ and FeCl₃ + O₂ reduced TP to 8 and 6%, respectively, of reference concentrations, whereas redox-insensitive treatments with alum or lime + alum reduced TP to 14% of reference concentrations. Lime and O₂ treatments reduced TP concentrations to 35 and 52% of reference concentrations, respectively.
4. The fraction of P that adsorbed and co-precipitated with iron and aluminium in the sediment cores was low (non-apatite phosphorus fractions < 5%), suggesting that P release was controlled by apatite solubility and bacterial metabolism.     

Chironomid stratigraphy in the shallow and eutrophic Lake Søbygaard, Denmark: chironomid–macrophyte co-occurrence

1. A sediment core from the shallow, hypertrophic Lake Søbygaard (mean depth ∼1 m; [TP] 310 μg P L⁻¹) was analysed for subfossil remains to reconstruct chironomid community changes in relation to the succession and disappearance of aquatic macrophytes. 2. Species composition in the 1.10 m core indicates a succession from a 'naturally' eutrophic state to a hypertrophic state during recent centuries. Radiometric dating (²¹⁰Pb) of the uppermost 20 cm of the sediment core (∼1932–93) indicates that sediment accumulation rate had doubled in recent decades. 3. Changes in chironomid assemblages were in close agreement with changes in both diatoms and macrophyte remains in the same core. Distinct changes in chironomid communities reflect the eutrophication process and macrophyte succession through Chara , Ceratophyllum and Potamogeton dominance to the present state, with complete loss of submerged vegetation and dominance by phytoplankton. 4. The co-occurrence and relationship between aquatic macrophyte diversity and recent subfossil chironomid assemblages were assessed from an additional 25 Danish lakes. There was good agreement between the macrophyte and chironomid-based lake groupings. Overall, a significant difference ( P <0.001) was found in chironomid assemblages among lakes in different macrophyte classes. In a pair-wise comparison, the poorly buffered mesotrophic lakes and the alkaline eutrophic lakes had significantly different chironomid assemblages. 5. Chironomid taxa commonly reported to be associated with macrophytes ( Cricotopus , Endochironomus and Glyptotendipes ) were shown also to be indicators of highly productive lakes lacking abundant submerged vegetation.     

Ceratophyllum demersum hampers phytoplankton development in some small Norwegian lakes over a wide range of phosphorus concentrations and geographical latitude

1. Data on submerged and floating-leafed macrophytes, phytoplankton, nutrients (N, P) and calcium were collected from twenty-four small lakes ( 1 km²) over a wide range of latitudes in Norway. The majority of the investigated lakes were mesotrophic or eutrophic, and most of the lakes were markedly affected by diffuse and point-source runoff from agriculture. According to their macrophyte species composition, the majority of the lakes can be classified as Potamogeton lakes or Chara lakes, or a combination of these.
2. This study is consistent with the 'two alternative stable states' hypothesis. We observed clearwater lakes with dense macrophyte cover over a wider range of total P concentration than has been reported previously: from 30 to more than 700 mg P m^–3. The clearwater state was only observed in lakes with mean depths of less than 1.9 m.
3. Most clear lakes with high cover of submerged vegetation showed indications of N limitation.
4. In this study nearly all the macrophyte-dominated lakes with P concentrations above 30 mg m^–3 had dense stands of Ceratophyllum demersum (L.). This indicates that Ceratophyllum may also play an important role in stabilizing and maintaining a clearwater state at high P concentrations.     

Ceratophyllum demersum hampers phytoplankton development in some small Norwegian lakes over a wide range of phosphorus concentrations and geographical latitude

1. Data on submerged and floating-leafed macrophytes, phytoplankton, nutrients (N, P) and calcium were collected from twenty-four small lakes ( 1 km²) over a wide range of latitudes in Norway. The majority of the investigated lakes were mesotrophic or eutrophic, and most of the lakes were markedly affected by diffuse and point-source runoff from agriculture. According to their macrophyte species composition, the majority of the lakes can be classified as Potamogeton lakes or Chara lakes, or a combination of these.
2. This study is consistent with the 'two alternative stable states' hypothesis. We observed clearwater lakes with dense macrophyte cover over a wider range of total P concentration than has been reported previously: from 30 to more than 700 mg P m^–3. The clearwater state was only observed in lakes with mean depths of less than 1.9 m.
3. Most clear lakes with high cover of submerged vegetation showed indications of N limitation.
4. In this study nearly all the macrophyte-dominated lakes with P concentrations above 30 mg m^–3 had dense stands of Ceratophyllum demersum (L.). This indicates that Ceratophyllum may also play an important role in stabilizing and maintaining a clearwater state at high P concentrations.     

Changes in abundance, composition and controls within the plankton of a fertilised arctic lake

1. An oligotrophic arctic lake was fertilised with inorganic nitrogen and phosphorus as (NH₄)₂ NO₃ and H₃PO₄ for five summers. The loading rate was 1.7–2.5 mmol N m^–2 day^–1 and 0.136–0.20 mmol P m^–2 day^–1 which is two to three times the annual loading of lakes in the area. The heterotrophic microzooplankton community was enumerated during the experiment as well as 1 year pre- and post-treatment.
2. The structure of the microplankton community changed from a nutrient limited system, dominated by oligotrich protozoans and small-particle feeding rotifers, to a system dominated by a succession of peritrich protozoans and predatory rotifers. These peritrich protozoans and predatory rotifers were not present prior to fertilisation and never constituted more than a small fraction of the biomass in other lakes at the research site. The average biomass of the rotifers and protozoans was more than seven and a half times larger by the end of fertilisation than it was initially.
3. Because of the increases in numbers of individuals in these new taxa, the structure of the microbial food web changed. When fertilisation stopped, most parameters returned to prefertilisation levels within 1 year.     

Polyaluminum chloride-enhanced concentration efficiency of poliovirus and f2 phage from sewage water

Aims:  To enhance the recovery of f₂ bacteriophage and poliovirus by an established method based on the adsorption to and elution from positively-charged Al(OH)₃-treated silica gel.
Methods and Results:  Polyaluminum Chloride (PAC) was added to water samples to neutralize the negatively charged materials, which can reduce virus recovery by providing a competing adsorption mode on the media surface. Using this improved process (PAC 30 mg l⁻¹, pH 6·5, temperature 20∼30°C), the recoveries of Poliovirus I and f₂ from small-volume sewage (100 ml) were 110·76 ± 36·0% and 92·06 ± 8·65%, respectively ( P  < 0·05 vs. traditional methods). Recovery from a 20-L volume of sewage averaged 85·65 ± 4·43% for f₂ and 88·73 ± 9·76% for poliovirus, significantly higher than the recoveries in the traditional methods ( P  < 0·05).
Conclusions:  PAC could enhance concentration efficiency of poliovirus and f₂ phage from sewage water.
Significance and Impact of the Study:  This method should significantly improve the recovery of viruses from sewage.     

The effect of an emergent macrophyte (Typha angustifolia) on sediment resuspension in a shallow north temperate lake

1. The effects of emergent macrophytes on water turbidity and sediment resuspension in the shallow Kirkkojärvi basin of Lake Hiidenvesi were studied with sediment traps, and concomitant sediment and water samples. The study was conducted during May–August in three different zones of a stand of the emergent Typha angustifolia .
2. Within the stand (5 m from the edge), both the concentration of suspended solids and the rate of sediment resuspension were significantly lower than at the edge and outside the stand (5 m from the edge). The differences between the zones increased towards the end of summer together with the growing stem density. During the study period (82 days), 2210 g dw m⁻² of sediment was resuspended in the outer zone. At the edge and in the inner zone, the corresponding numbers were 1414 and 858 g dw m⁻², respectively.
3. With the resuspended sediment, 39.4 mg P  m⁻² day⁻¹ was brought to the water column outside the stand, 22.4 mg P  m⁻² day⁻¹ at the edge and 13.4 mg P  m⁻² day⁻¹ within the stand.
4. In early summer, the concentration of suspended solids had a highly significant positive effect on soluble reactive phosphorus (SRP) concentration in the water, whereas in late summer no effect was found. During the study period, phosphorus retention by emergent macrophyte stands corresponded to 3–5% of the present annual external phosphorus loading of the Kirkkojärvi basin.     

Measurements of phosphorus uptake by macrophytes and epiphytes from the LaPlatte River (VT) using 32P in stream microcosms

1. Phosphorus (P) uptake by macrophytes and epiphytes from the LaPlatte River (VT) was examined in the laboratory by adding ³²PO₄‐P to recirculating stream microcosms.
2. Water, plugs of sediment and plants were removed from the river and placed into the microcosms. ³²PO₄‐P was then added either to the water or the sediment, and its incorporation into plants and epiphytes was monitored over 3 days. Uptake was examined at both ambient (5 μg L^–1) and increased (50 μg L^–1) soluble reactive phosphorus (SRP) concentrations. A computer program was developed to fit curves to the radiotracer data and calculate rate constants for the simultaneous transfer of ³²P among compartments.
3. Both macrophytes and epiphytes removed P from the water, but epiphyte uptake of P was more rapid. Phosphate enrichment stimulated P uptake by both macrophytes and epiphytes. Macrophytes also obtained P from the sediment. The relative contribution of P to macrophytes from the water vs. that from the sediment appeared to vary with SRP in the overlying water. Accurate estimates of rates of P uptake from sediments by macrophytes were difficult to obtain however, due to very low and highly variable unit rate constants for P uptake and uncertainty about the magnitude of the phosphate pool available for uptake.
4. SRP concentrations were greater in the overlying water than in the sediment pore water of stream microcosms in the present study. Numerous reports in the literature have suggested that this condition favours uptake by macrophyte stems and leaves rather than by roots.
5. Phosphate uptake from the water by macrophytes in shallow streams may be more common than for macrophytes in lakes.     

Lough Ennell: laboratory studies on sediment phosphorus release under varying mixing, aerobic and anaerobic conditions

SUMMARY. 1. The exchange of phosphorus between the epilimnetic (shallow zone) sediment and water column in Lough Ennell was investigated in laboratory experiments using five intact cores.
2. Variations in water mixing, sediment suspension and aerobic–anaerobic oxygen status in the water column and its effects on sediment phosphorus release rates were determined.
3. Experimental results indicated that phosphorus release is possible under both aerobic and anaerobic conditions. Aerobic release (0.025 mg P l⁻¹ over 5 days) was possible up to the point when mass resuspension of sediment occurred. Anaerobic release for the same period and mixing conditions was 0.183 mg Pl⁻¹.
4. The release rate under aerobic conditions at 10°C equates to an internal areal loading of 0.134 g P m⁻² yr^−1, which is approximately 17% and 30% of the average total phosphorus and orthophosphate loadings respectively for the period 1974–79.
5. The results clearly implicate aerobic inorganic phosphorus release from the epilimnetic sediments as a significant source of this nutrient to the overlying water column and is likely a major factor in the continuing eutrophic status in the lake.     

Effects of CO2 enrichment on mineral accumulation and nitrogen relations in a submersed macrophyte

1. Six- to eight-week greenhouse experiments with independent control of pH and dissolved CO₂ evaluated the potential for CO₂ enrichment to stimulate the accumulation of Al, Fe, P and N in shoots of Vallisneria americana , particularly at pH 5. These minerals were provided only as they occurred in natural lake sediments.
2. The effect of CO₂ enrichment at pH 5 v pH 7.3 on growth and tissue N concentration was also determined.
3. CO₂ enrichment at pH 5 effected 5.5- and 7-fold increases in total shoot accumulation of Al and Fe, respectively. In a two-way factorial experiment, CO₂ enrichment yielded 6- to 11-fold greater total shoot P accumulation in plants grown on less and more fertile sediments, respectively.
4. In a three-way factorial experiment, CO₂ enrichment stimulated Vallisneria growth, especially at pH 5, and resulted in a 31–58% reduction in tissue [N] for different pH × sediment combinations. These are greater reductions than previously reported. It also increased total shoot N accumulation up to 6-fold, and there were significant interactions with pH and sediment source: the CO₂ enrichment effect on shoot N accumulation was greater at pH 5 than at pH 7.3, and it was greater with the more fertile sediment at pH 5.
5. Water chemistry (pH and/or [CO₂]) and sediment fertility thus both indirectly influenced the accumulation of sediment-derived minerals in macrophyte shoots within the water column.     

Effects of CO2 enrichment on mineral accumulation and nitrogen relations in a submersed macrophyte

1. Six- to eight-week greenhouse experiments with independent control of pH and dissolved CO₂ evaluated the potential for CO₂ enrichment to stimulate the accumulation of Al, Fe, P and N in shoots of Vallisneria americana , particularly at pH 5. These minerals were provided only as they occurred in natural lake sediments.
2. The effect of CO₂ enrichment at pH 5 v pH 7.3 on growth and tissue N concentration was also determined.
3. CO₂ enrichment at pH 5 effected 5.5- and 7-fold increases in total shoot accumulation of Al and Fe, respectively. In a two-way factorial experiment, CO₂ enrichment yielded 6- to 11-fold greater total shoot P accumulation in plants grown on less and more fertile sediments, respectively.
4. In a three-way factorial experiment, CO₂ enrichment stimulated Vallisneria growth, especially at pH 5, and resulted in a 31–58% reduction in tissue [N] for different pH × sediment combinations. These are greater reductions than previously reported. It also increased total shoot N accumulation up to 6-fold, and there were significant interactions with pH and sediment source: the CO₂ enrichment effect on shoot N accumulation was greater at pH 5 than at pH 7.3, and it was greater with the more fertile sediment at pH 5.
5. Water chemistry (pH and/or [CO₂]) and sediment fertility thus both indirectly influenced the accumulation of sediment-derived minerals in macrophyte shoots within the water column.     

Invertebrate grazing during the regenerative phase affects the ultimate structure of macrophyte communities

1. Although the biomass of freshwater macrophytes consumed by invertebrate herbivores (excluding crayfish) is usually low, there is growing evidence that invertebrates do exert a structuring effect on macrophyte communities. To explain this, we postulated that the effect of invertebrates may be concentrated on macrophytes during their regenerative phase.
2. We tested this hypothesis by means of a mesocosm-based experiment, in which we investigated the effects of different densities of pond snails [ Lymnaea stagnalis (L.)] on macrophytes regenerating from the natural sediment propagule bank.
3. After 2 months, a diverse macrophyte community had established in the absence of snails, mainly from sexual propagules. Under moderate snail grazing (4 individuals m⁻²), the ultimate biomass of macrophytes was similar, but its species composition differed dramatically. Only a few unpalatable taxa, such as Ceratophyllum demersum and Nymphaeaceae, persisted. Moreover, the relative success of macrophytes regenerating from vegetative rather than sexual propagules improved. Under higher snail grazing (20 m⁻²), all macrophytes disappeared before the end of the experiment.
4. These results confirm that snails at natural densities can have a strong effect on the ultimate structure of macrophyte communities by selectively consuming some species at a juvenile stage. Therefore, the regenerative phase can be seen as a window of opportunity for invertebrate grazers, which can have a qualitative effect on communities that is disproportionate to the biomass consumed.     

Taxonomic composition and biomass of heterotrophic flagellates in relation to lake trophy and season

1. Taxonomic composition and abundance of heterotrophic flagellates (HF) were studied in 55 lakes with different trophy in northern Germany using a live-counting technique.
2. Mean abundances and biomasses of HF ranged from 169 cells L⁻¹ and 22 μg L⁻¹ in mesotrophic lakes to 2439 cells mL^–1 and 475 μg L⁻¹ in hypertrophic lakes, respectively. Highest values were generally observed in spring, but mesotrophic lakes showed maximum values in early summer.
3. The taxonomic composition of HF was not significantly influenced by lake trophy and season. The major fraction of HF consisted of chrysomonads and Protista incertae sedis; other important groups were choanoflagellates and bicosoecids. The size distribution of HF changed with lake trophy and season, with a higher proportion of large HF (> 10 μm) in hypertrophic lakes and in spring, respectively.
4. Correlation analyses revealed a strong negative impact of cladocerans on total HF biomass and especially on large HF. Ciliates and large bacteria (> 10 μm) were strongly positively correlated with HF biomass; small bacteria (< 2 μm) showed a weak positive correlation.
5. Analyses at the level of species and genera revealed distinct distribution patterns of some taxa. Paraphysomonas , Aulacomonas and Quadricilia as large-bodied HF showed highest abundance in hypertrophic lakes and in spring. Attached taxa (e.g. Monosiga , Salpingoeca amphoridium ) were highly abundant in late summer and autumn, whereas Spumella and Kathablepharis occurred frequently in most samples.     

Mineralization, pore water chemistry and phosphorus release from peaty sediments in the eutrophic Loosdrecht lakes, The Netherlands

SUMMARY. 1. Pore water chemistry in peaty sediment was monitored for a year at two representative locations of the eutrophic shallow Loosdrecht lakes. The Netherlands. Phosphorus fluxes over the sediment-water interface were calculated using measured concentration gradients in the pore water and compared to fluxes measured under laboratory conditions. Results were analysed with Redundancy Analysis to detect patterns of variation in pore water chemistry and in measured and calculated fluxes, that could be ascribed to environmental variables.
2. It was demonstrated that phosphorus fluxes measured in long-term laboratory incubations were not correlated to any of the pore water characteristics.
3. Initial phosphorus fluxes measured in sediment columns, which varied between −7.7 and 1330 μmol m⁻²: day⁻¹, were correlated significantly to the calculated phosphorus flux over the sediment-water interface.
4. The high correlation between calculated fluxes of ammonia, phosphorus and methane and measured initial flux of phosphorus, conclusively pointed to mineralization of organic matter as the driving force for phosphorus release from the sediment.
5. Redundancy Analysis demonstrated that the rates of mineralization and phosphorus release were only weakly related to temperature. They appeared to be especially stimulated by the autumnal decrease in temperature which was probably related to an extra input of organic matter.     

The nutrient budget of a small eutrophic loch and the effectiveness of straw bales in controlling algal blooms

1. A 1-year intensive study of nutrient flows in Airthrey Loch, central Scotland, a small eutrophic [mean total phosphorus (TP) = 61.1 μg P l^–1] well-flushed freshwater body (area, 6.9 ha; mean depth, 1.85 m; volume, 1.274 × 10⁵ m³; retention time, 0.44 yr), was undertaken.
2. The nutrient budget was dominated by large allochthonous P inputs, equivalent to an areal load to the loch of 8.56 kg P ha^–1 yr^–1, which occurred predominantly during winter. In the summer, when TP inputs were low, water column levels of TP still increased, as a result of aerobic P release from sediments.
3. Sorption experiments indicated potential for sediment P release at water P concentrations of up to 200 μg P l^–1.
4. Aerobic release rate of P from sediments to the water column of Airthrey Loch was estimated to be of the order of 1 mg P m^–2 day^–1, and occurred during periods of elevated water column pH.
5. Straw bales placed in the loch to retard algal blooms were found not to have any demonstrable impact on algal concentrations observed during the study.     

The nutrient budget of a small eutrophic loch and the effectiveness of straw bales in controlling algal blooms

1. A 1-year intensive study of nutrient flows in Airthrey Loch, central Scotland, a small eutrophic [mean total phosphorus (TP) = 61.1 μg P l^–1] well-flushed freshwater body (area, 6.9 ha; mean depth, 1.85 m; volume, 1.274 × 10⁵ m³; retention time, 0.44 yr), was undertaken.
2. The nutrient budget was dominated by large allochthonous P inputs, equivalent to an areal load to the loch of 8.56 kg P ha^–1 yr^–1, which occurred predominantly during winter. In the summer, when TP inputs were low, water column levels of TP still increased, as a result of aerobic P release from sediments.
3. Sorption experiments indicated potential for sediment P release at water P concentrations of up to 200 μg P l^–1.
4. Aerobic release rate of P from sediments to the water column of Airthrey Loch was estimated to be of the order of 1 mg P m^–2 day^–1, and occurred during periods of elevated water column pH.
5. Straw bales placed in the loch to retard algal blooms were found not to have any demonstrable impact on algal concentrations observed during the study.     

Population dynamics and production of Jesogammarus annandalei, an endemic amphipod, in Lake Biwa, Japan

SUMMARY 1. Population dynamics and production of Jesogammarus annandalei , an endemic amphipod in Lake Biwa, were examined from April 1997 to June 1998. The life cycle of this species was 1 year with the new generation beginning in early autumn. They preferred low temperature (<12 °C) and their spatial distribution varied seasonally and accordingly.
2. In deep water, the abundance of J. annandalei ranged from 200 to 63 000 m⁻² and decreased towards summer and the biomass (0.01∼3.6 g C m⁻²) was on average comparable that of zooplankton. The density was much higher than that recorded by a study conducted 35 years ago.
3. Individual growth rate of this amphipod was high in winter and spring but decreased in summer. Annual production of J. annandalei (6.2 g C m⁻² year⁻¹) was only 2% of primary production but was at the higher end of the range reported for amphipods in oligo- and mesotrophic lakes.
4. These results are consistent with the view that Lake Biwa is becoming more eutrophic, with a consequent decrease in the abundance of predatory fish in the profundal zone.