Dynamics and ecological significance of daily internal load of phosphorus in shallow Lake Balaton, Hungary

1. As supported by field data, turbidity recorded by light scattering sensors could reliably be converted into concentration of suspended particulate matter (SPM) and coefficient of vertical light attenuation (K_d) in Lake Balaton. 2. Autocorrelation analysis revealed that proper determination of SPM concentration and K_d required daily sampling. To approximate daily rate of resuspension, 15 min or more frequent measurements were needed. Thus, routine monitoring provides very little insight into environmental variability of shallow lakes as habitats for phytoplankton. 3. The internal P load was estimated from daily rate of resuspension and P desorption capacity of sediments. The latter was assumed to be proportionate to the potentially mobile inorganic P content of SPM. A comparison with net primary production and nutrient status of phytoplankton showed that the proposed method of estimating time series of internal P load captured seasonal trends. 4. The daily rate of resuspension was high whereas that of internal P load was low in Lake Balaton relative to other shallow lakes. The latter reflects favourable behaviour of the calcite‐rich sediments. As a consequence, carrying capacity of Basin 1 of Lake Balaton was P‐determined. 5. The timing of external and internal loads was radically different. While the former showed mostly seasonal changes, large pulses characterised the latter. As a consequence, internal load may supply more P to phytoplankton growth during the critical summer months than external load. However, the relative importance of these sources may show substantial interannual variability. 6. Large resuspension events often followed each other during periods of 10–15 days. It has been shown that disturbances in this frequency range are of key importance in maintaining the diversity of phytoplankton. We propose that resuspension can be perceived not only as a disturbance factor but also as a factor that periodically relaxes nutrient stress. The former feature may dominate the instantaneous effect, whereas the latter may determine the persistent effect of resuspension on succession of phytoplankton.     

滇池、红枫湖沉积物中总磷、分态磷及生物硅形态与分布特征

通过对不同富营养化程度的湖泊（滇池及红枫湖）沉积柱剖面进行不同形态磷以及生物硅含量分析,对比了磷及生物硅的沉积分布特征,并探讨了其与湖泊富营养化演化的关系。结果表明：滇池沉积物中碎屑磷和有机磷在总磷中占50.5%～80.8%,是影响总磷趋势的主要原因;红枫湖中有机磷对总磷的贡献较大,占总磷的53%～60.9%;且两湖各形态磷均随深度的增加而减少;两湖总磷、有机磷、铝磷以及滇池沉积柱中的碎屑磷等的快速增长,表明其主要来源于人为污染;生物硅含量呈现从下层沉积物向上逐渐增加,在上层沉积物中逐渐减少的趋势;总磷、有机磷、铁结合态磷、铝结合态磷、弱结合态磷以及生物硅都能较好地反映富营养化污染的演变和现状。     

Nitrogen dynamics in Lake Okeechobee: forms,functions, and changes

Total nitrogen (TN) in Lake Okeechobee, a large, shallow, turbid lake in south Florida, has averaged between 90 and 150 μM on an annual basis since 1983. No TN trends are evident, despite major storm events, droughts, and nutrient management changes in the watershed. To understand the relative stability of TN, this study evaluates nitrogen (N) dynamics at three temporal/spatial levels: (1) annual whole lake N budgets, (2) monthly in-lake water quality measurements in offshore and nearshore areas, and (3) isotope addition experiments lasting 3 days and using ¹⁵N-ammonium (¹⁵NH₄ ⁺) and ¹⁵N-nitrate (¹⁵NO₃ ⁻) at two offshore locations. Budgets indicate that the lake is a net sink for N. TN concentrations were less variable than net N loads, suggesting that in-lake processes moderate these net loads. Monthly NO₃ ⁻ concentrations were higher in the offshore area and higher in winter for both offshore and nearshore areas. Negative relationships between the percentage of samples classified as algal blooms (defined as chlorophyll a > 40 μg l⁻¹) and inorganic N concentrations suggest N-limitation. Continuous-flow experiments over intact sediment cores measured net fluxes (μmol N m⁻² h⁻¹) between 0 and 25 released from sediments for NH₄ ⁺, 0–60 removed by sediments for NO₃ ⁻, and 63–68 transformed by denitrification. Uptake rates in the water column (μmol N m⁻² h⁻¹) determined by isotope dilution experiments and normalized for water depth were 1,090–1,970 for NH₄ ⁺ and 59–119 for NO₃ ⁻. These fluxes are similar to previously reported results. Our work suggests that external N inputs are balanced in Lake Okeechobee by denitrification.     

Excretion of total phosphorus,dissolved reactive phosphorus,ammonia, and urea by Lake MichiganMysis relicta

Rates of nutrient release byMysis relicta in Lake Michigan were measured on five nights at a 45-m station near Milwaukee, Wisconsin, U.S.A., in the summer of 1977. Nocturnal vertical migrations of the mysids were monitored with both echosounder tracings and vertical net tows. Estimates of the total areal dry mass of the mysids ranged from 600 to 1 820 mg m–2. Rates of release of dissolved reactive phosphorus, total phosphorus, ammonia, and urea were measured in dark incubations on shipboard. Excretion experiments were initiated immediately after mysids were collected from each of several vertical net hauls. The depths of maximum mysid densities corresponded approximately with a deep phytoplankton peak located in the vicinity of the thermocline. Semiquantitative demands for N and P by phytoplankton within this peak were obtained from¹⁴C estimates of primary production from a previous study, assuming a constant C:N:P ratio for the algae. These algal nutrient demands were compared to potential N and P release by the mysids to obtain a first approximation of the relative rates of nutrient supply and demand for the field phytoplankton populations. Our analysis indicates that mysids may directly supply about 1–10% of the daily N and P demands of the phytoplankton in the deep peak. However, indirect interactions betweenMysis relicta and other organisms, such as small zooplankton and fishes, could be major factors in nutrient recycling within the metalimnion and subthermocline region of Lake Michigan.Contribution No. 238, Center for Great Lakes Studies, University of Wisconsin-Milwaukee, Wisconsin.Contribution No. 238, Center for Great Lakes Studies, University of Wisconsin-Milwaukee, Wisconsin.     

Biotic feedbacks in Lake phosphorus cycles

Limnologists are now reconsidering the role of the biota in the phosphorus (P) cycles of lakes. Changes in lake communities can have significant consequences for ecosystem P cycles. At seasonal timescales, the relative importance of nitrogen (N) and Pas limiting factors for primary production depends in part on zooplankton species composition. Phosphorus storage and recycling by fish and zooplankton can be large components of P budgets, and mobile consumers can be important vectors in P transport. Stability, resilience and resistance of lake P cycles may depend heavily on fluxes to and from upper trophic levels.     

The influence of water circulation on chlorophyll-turbidity relationships in Lake Okeechobee as determined by remote sensing

The spatial distribution of phytoplankton can be difficult toassess in shallow, productive aquatic systems due to frequentalgal blooms, high turbidity and sediment-resuspension events.We conducted a study to assess the distribution of suspendedparticles in Lake Okeechobee, Florida, utilizing both Landsat(1974–75) or Advanced Very High Resolution Radiometer(AVHRR) (1987) satellite remote sensing. Surface water sampleswere collected by helicopter to determine in situ chlorophyll-aand turbidity levels at 20 stations on four dates in 1974–75and six dates in 1987. Remotely sensed reflectance values agreedwell with in situ particle densities at the 20 in-lake stations(average R²: Landsat = 0.81, AVHRR = 0.53) and independent,synoptic boat mapping of algal blooms (r² = 0.79, P < 0.01).Basin-wide maps of chlorophyll and turbidity, as well as additionalspatial sampling, both indicated that these parameters are notnecessarily coupled in Lake. Our data concur with the hypothesisthat the spatial distributions of chlorophyll and turbidityare shaped by different forces. The highest concentrations ofchlorophyll occurred in the vicinity of tributary nutrient inputsat the lake's perimeter, while turbidity increased towards thecenter of the lake, reflecting predominant water circulationpatterns. ²Present address: Department of Biology & Romberg TiburonEnvironmental Center, San Francisco State University, San Francisco,CA 94132, USA ³Present address: Idaho Division of Environmental Quality, 1420North Hilton, Boise, ID 83706-1260, USA ⁴Present address: 5642 Santiago Circle, Boca Raton, FL 33433,USA     

Water quality effectiveness of ditch fencing and culvert crossing in the Lake Okeechobee basin, southern Florida, USA

Ditch fencing and culvert cattle crossing Best Management Practice (BMP) was evaluated in this study with regard to phosphorus (P) and nitrogen (N) load reductions and economic feasibility in the Lake Okeechobee (LO) basin. The BMP was implemented at a 170 m section of a drainage ditch within a ranch in the LO basin and flow and concentration (N and P) data at the upstream and downstream of the ditch were collected for one pre-BMP (June-October, 2005) and three post-BMP (June-October, 2006-2008) periods. During the pre-BMP period, downstream total P (TP) load was 20% (67.0 kg) higher than the upstream, indicating the cattle crossing to be a source of P. Downstream loads of TP in 2006 and 2008 (post-BMP periods) became 26% (14.7 kg) and 11% (85.9 kg) lower than the upstream loads, respectively indicating that the BMP reduced the P loads. The site was a sink for N for all periods except the 2007. Unusual dry conditions during 2007 resulted in the addition of P and N at the BMP site, probably due to the release of P and N from soil and plants. Average of three post-BMP period load showed a 10% reduction of TP loads at the downstream (251.8 kg) compared to the upstream (281.0 kg) location. To consider potential P contributions from the soil and plant, two scenarios, conservative and liberal, were considered to estimate P load reductions due to the BMP. For the conservative scenario, P contribution from soil and plant was considered, while for liberal it was not. Reductions in P loads for conservative and liberal scenarios were 0.35 and 0.44 kg/day, respectively. Phosphorus removal cost for the conservative scenario was $12.61/kg of P, which is considerably less than the cost of other P reduction strategies in the basin. Overall, results show that the BMP can reduce P concentration and loads from ranches without causing adverse impact on cattle production.     

Revisiting the surgical creation of volume load by aorto-caval shunt in rats

Cardiac hypertrophy is an early landmark during the clinical course of heart failure, and is an important risk factor for subsequent morbidity and mortality. The hypertrophy response to different types of cardiac overload is distinguished both at the molecular and cellular levels. These changes have been extensively characterized for pressure load hypertrophy; however, similar information for volume load hypertrophy is still needed. This study was undertaken to improve the existing method of producing experimental cardiac volume load. Previous investigators have employed surgical aorto-caval shunt (ACS) as a model for volume load hypertrophy (VO) in rats. The procedure is relatively simple and involves glue to seal the aortic hole after ACS. However, it has several limitations mostly related to the use of glue e.g. poor visualization due to hardening of tissues, imperfect sealing of the puncture site and glue seeping through the aortic hole resulting in shunt occlusion. We have modified the procedure using aortic adventitial suture instead of glue and 18G angiocatheter instead of 16G needle, which eliminated the technical difficulties from the former method. The ACS was visually confirmed at sacrifice, and the VO demonstrated by time-related changes in the heart weight/body weight ratio which increased from 78% at 4 weeks to 87% at 10 weeks and increased liver/body weight ratio by 22% at 10 weeks of post aorto-caval shunt. Cardiac expression of atrial natriuretic peptide (ANF) also demonstrated time-related increase in ANF mRNA (+275% increase at 4 weeks, p < 0.05, and +370% increase at 10 weeks, p < 0.001). This modified technique of aorto-caval shunt offers simpler, reproducible and consistent model for VO hypertrophy in rats.     

Summer dynamics of the deep chlorophyll maximum in Lake Tahoe

Vertical profiles of chlorophyll and phytoplankton biomass weremeasured in Lake Tahoe from July 1976 through April 1977. Adeep chlorophyll maximum (DCM) persisted during summer and earlyautumn (July—October) near 100 m, well below the mixedlayer and at the upper surface of the nitracline. The DCM coincidedwith the phytoplankton biomass maximum as determined from cellcounts. In addition, the composition of the phytoplankton assemblagewas highly differentiated with respect to depth. Cyclotellastelligera was the predominant species in the mixed layer whilethe major species in the DCM layer included C. ocellata andseveral green ultraplanktonic species. In situ cell growth playsa substantial role in maintaining the DCM, but sinking of cellsfrom shallower depths and zooplankton grazing above the DCMmay contribute to the maintenance of the DCM. Calculations supportthe interpretation that the summer DCM persists at the boundarybetween an upper, nutrient-limited phytoplankton assemblageand a deeper, light-limited assemblage.     

Seasonal and spatial variation in zooplankton community structure and their relation to possible controlling variables in Lake Okeechobee

1. We examined the behavioural response of stream macroinvertebrates to real and simulated predatory activity by a river bird, the Eurasian dipper, Cinclus cinclus L.
2. In the field, we assessed whether invertebrate drift changed in response to live dippers in enclosures; we found no effects on the drift of any of the five families for which individual analyses were possible, either because it was infrequent, or involved distances too short to be detected (< 0.5 m).
3. In a laboratory stream, we observed prey during encounters with a model dipper which simulated flight, swimming, bill contact with the prey, and stone turning. Invertebrate families varied in their response. Simuliids and hydropsychid caddis lacked effective escape behaviour, consistent with heavy losses through predation by dippers in the wild. Other families either drifted (Baetidae, Gammaridae) or moved away (Heptageniidae, Ephemerellidae, Leuctridae, Perlidae) from the model dipper, but responded only to bill contact or simulated stone turning. Such delayed responses would not protect individuals directly targeted by foraging dippers and partly explain the lack of detectable effects by dippers on drift in the field.
4. We suggest why invertebrates do not show more marked escape responses to this important predator.     

Seasonal and spatial variation in zooplankton community structure and their relation to possible controlling variables in Lake Okeechobee

1. We examined the behavioural response of stream macroinvertebrates to real and simulated predatory activity by a river bird, the Eurasian dipper, Cinclus cinclus L.
2. In the field, we assessed whether invertebrate drift changed in response to live dippers in enclosures; we found no effects on the drift of any of the five families for which individual analyses were possible, either because it was infrequent, or involved distances too short to be detected (< 0.5 m).
3. In a laboratory stream, we observed prey during encounters with a model dipper which simulated flight, swimming, bill contact with the prey, and stone turning. Invertebrate families varied in their response. Simuliids and hydropsychid caddis lacked effective escape behaviour, consistent with heavy losses through predation by dippers in the wild. Other families either drifted (Baetidae, Gammaridae) or moved away (Heptageniidae, Ephemerellidae, Leuctridae, Perlidae) from the model dipper, but responded only to bill contact or simulated stone turning. Such delayed responses would not protect individuals directly targeted by foraging dippers and partly explain the lack of detectable effects by dippers on drift in the field.
4. We suggest why invertebrates do not show more marked escape responses to this important predator.     

Mobility of phosphorus fractions in the sediments of Lake Balaton

Sediment phosporus was studied by a combination of the fractionation procedure of Hieltjes & Lijklema and the isotopic dilution technique in a mesotrophic (Tihany) and a hypertrophic (Keszthely) basin of Lake Balaton.In the calcareous sediments the largest part of phosphorus was bound to calcium. Iron-bound and residual P showed higher concentrations at Keszthely than at Tihany. There was little loosely adsorbed P at both locations. Vertical differences in P fractions of the sediments were more pronounced at Keszthely than at Tihany. Exchangeability of the fractions decreased in the following sequence: iron-bound > loosely adsorbed > calcium-bound > residual. Phosphorus, particularly in the calcium-bound fraction, was more mobile at Keszthely, and its exchangeability decreased rapidly downwards at both stations. Four times more potentially mobile phosphorus has been accumulated by the upper sediment layer at Keszthely than at Tihany.     

Periphyton nutrient limitation and other potential growth-controlling factors in Lake Okeechobee,U.S.A.

Periphyton nutrient limitation was assessed in Lake Okeechobee, a large, shallow, eutrophic lake in the southeastern U.S.A. Nutrient assays were performed to determine if the same nutrients that limit phytoplankton also limit periphyton growth in the lake. Nutrient diffusing clay substrates containing agar spiked with nitrogen, phosphorus, or both, along with nutrient-free controls, were incubated at four sites in the lake. Three sites were located in a pelagic–littoral interface (ecotone) and one site was located in the interior littoral region. Incubations lasted for 20–26 days, and were repeated on a quarterly basis between 1996 and 1997, to incorporate seasonal variability into the experimental design. The physical and chemical conditions at each site also were measured. Periphyton biomass (chlorophyll a and ash-free dry mass) was highest at the littoral and northern ecotone sites. At the littoral site, nitrogen limited biomass in four of five incubations, although the largest biomass differences between the treatments and controls (3 g cm^–2 as chl) were probably not ecologically significant. Periphyton biomass at the western and southern ecotone sites was low compared to the other two sites. Increases in water column depth and associated declines in light penetration strongly correlated with periphyton growth and suggested that they may have limited growth most often at all three ecotone sites. Nitrogen also was found to limit periphyton growth approximately 20% of the time at the ecotone sites and phosphorus was found to limit growth once at the west site.     

Aquatic vegetation and largemouth bass population responses to water-level variations in Lake Okeechobee,Florida (USA)

A five-year study examined the responses of submerged aquatic vegetation (SAV), emergent vegetation, and largemouth bass (Micropterus salmoides) to variations in water level in a large lake in Florida, USA. SAV was assessed using a combined transect survey/spatial mapping approach, emergent vegetation was quantified with aerial photography and GIS, and bass were surveyed by electro-shocking. During the period leading up to this study (1995–1999), water levels were high in the lake, and the SAV was reduced in spatial extent and biomass, compared to its condition in the early 1990s. Spatial extent of emergent vegetation also was low, and largemouth bass surveys indicated low densities and failure to recruit young fish into the population. This was attributed to the lack of critical vegetative habitat. In spring 2000, the lake was lowered by discharging water from major outlets, and this was followed by a regional drought. Water levels dropped by 1m, and there was widespread development of Chara lawns in shoreline areas, with coincident increases in water clarity. There was some germination of vascular SAV, but Chara was the extreme dominant, such that structural complexity remained low. There was no substantive improvement in bass recruitment. During 2001, water levels declined further, and emergent plants germinated in exposed areas of the lake bottom. SAV was restricted to sites farther offshore, and continued to be dominated by Chara. There again was no bass response. In 2002, conditions changed when water levels increased to a moderate depth, flooding shoreline habitat to 0.5m. Vascular SAV increased in biomass and spatial extent, such that the community developed a high structural complexity. At the same time, emergent aquatic plants developed dense stands along the western shoreline. Largemouth bass had a strong recruitment of young fish for the first time in 5years. Recruitment continued to be successful in 2003, when spatial extent of SAV was somewhat reduced by higher water but total biomass and diversity remained high. These results demonstrate an important effect of inter-annual variation in water depth on the population dynamics of aquatic plants and fish in a subtropical lake.     

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     

Determination of total phosphorus in lake sediments

Several methods are in use to measure total phosphorus in lake sediments making cross-study comparisons tenuous, and may lead to potential errors in calculating other phosphorus fractions. Four methods in common use in the limnological literature were compared using sediments spanning a range of organic content from 2 to 35% C. No significant differences were found in within-lake comparisons, although a method using persulfate oxidation was highly variable, and this variability was correlated with organic content suggesting inconsistent oxidation. The other methods (combustion, acid digestion with HClO₄, H₂SO₄ + H₂O₂) gave uniformly small variances.     

Response of lakes to the reduction of phosphorus load

On occasion of the publication of a book (H. SAS, ed.) a discussion of the predictability of the response of eutrofied lakes on a reduced external phosphorus loading is presented. Whereas the OECD study examines the correlations between nutrient loading and indicators for eutrofication for groups of lakes, the book discusses the direction of the response on reduction of the phosphorus loading for individual lakes. Although this is a valuable analysis that can increase the insight in the mechanisms controlling lake responses, neither the OECD study nor this book yield sufficiently unquestioned data that can be used directly in a a management framework. The relationships between loading and inlake concentration and between phosphate concentration and chlorophyll are discussed in some detail.     

A tentative recommendation for the maximum daily intake of selenium.

In order to make a tentative recommendation for the maximum acceptable daily intake of selenium, relevant data were compiled from the available literature. Normal daily intake of selenium from foods was estimated as about 100 mug, half of which comes from fish and shellfish (in an average adult Japanese). Intake of selenium from other sources was negligible. The amount of selenium excreted in the urine was found to be compatible with the estimated value of the daily oral intake. The range of the margin of safety was then estimated as 10 to 200 times the normal level on the basis of human and animal toxicity data. The variation of dietary selenium intake in the general population is discussed, leading to the conclusion that the consumers of large amounts of fish may ingest as much as 500 mug daily. Consequently, a value of 500 mug is proposed as the tentative maximum acceptable daily intake of selenium for the protection of human health.     

Lake restoration by reducing external phosphorus loading: the influence of sediment phosphorus release

SUMMARY. 1. This review considers the factors which determine the recovery of eutrophic lakes following a reduction in the external phosphorus loading.
2 The mean phosphorus content of a lake should decrease roughly in proportion to the reduction in phosphorus input. Where the lake phosphorus concentration does not decrease as predicted, then the release of phosphorus from the sediment is implicated.
3. The current understanding of the processes by which sediment phosphorus is mobilized and transported into the photic zone is described. The extent to which phosphorus release can maintain lake phosphorus concentrations following the reduction in external loading is influenced by: lake morphometry, flushing rate, sediment type, trophic state and history of enrichment.
4. A reduction in the phytoplankton biomass of a lake is dependent upon the size of the decrease in lake phosphorus concentration and the degree to which phosphorus limits primary production. The importance of phosphorus in limiting phytoplankton production tends to decrease with increasing lake trophic status.
5. Improvements in the condition of highly eutrophic lakes require very large reductions in external phosphorus loading, whereas in mildly enriched lakes moderate changes in the supply of phosphorus have noticeable effects on phytoplankton biomass.     

Phytoplankton dynamics within the subsurface chlorophyll maximum of Lake Michigan

The Lake Michigan deep chlorophyll maximum during July 1977was a broad band of phytoplankton –15 m thick rather thanthe thin layers observed in smaller lakes. The maximum concentrationof chlorophyll was typically four times surface values. ¹⁴Cuptake incubations showed that the deep plankton layer was aviable component of the pelagic ecosystem accounting for upto 60% of primary production on an areal (m²) basis. The LakeMichigan chlorophyll maximum was composed primarily of largediatoms, filaxnentous green algae, and flagellates. Althoughlarge scale physical processes such as seiches had some effecton the chlorophyll maximum, the basic integrity of the layerwas maintained by biological processes. In particular, epilinmeticnutrient depletion was apparently a major factor affecting thevertical distribution of phytoplankton. Application of a diffusionmodel to the chemical and chlorophyll data indicated that nutrientuptake and primary production by phytoplankton were more importantin determining chemical and chlorophyll vertical profiles thandiffusivity. ¹Contribution No. 389 from the Great Lakes Research Division,University of Michigan.