A comparison of chlorophyll a and carotenoid concentrations as indicators of algal volume

SUMMARY. 1. Carotenoid concentration, as measured hy absorbance at 480 nm. was a better indicator of algal volume than chlorophyll a when the results from two lakes and laboratory studies on Osciltaloria agardhii var. isothrix Skuja were compared. The correlation between algal volume and carotenoid in White Lough ( r =0.91) was significantly higher (0.001 P <0.01) than ihat between algal volume and chlorophyll a (r=0.77). The Lotigh Neagh correlation coefficient lor algal volume with carotenoid (r=0.89) was only marginally stronger than that with chlorophyll a (r=0.87).
2. The relatively weak correlation between algal volume and chlorophyll a in White Lough was a result of a summer decline in the chlorophyll a content of O. agardhii var. isothrix , which dominated the phytoplankton. The chlorophyll a content of the phytoplankton was depressed by high summer daily totals ol light hours received by the phytoplankton in White Lough of up to 14 h in comparison to a maximum value of 3.8 h in Lough Neagh. where no seasonal cycle of chlorophyll a content was evident.
3. Laboratory studies demonstrated that while chlorophyll a per unit algal volume of O. agardhii var. isothri.x declined with increasing light dose, carotenoid content did not. Nitrogen and phosphorus limitation depressed the carotenoid content but to a lesser degree than was observed for chlorophyll a.     

Decomposition of aquatic biota and sediment formation: lipid components of two blue-green algal species and of detritus resulting from microbial attack

To simulate early stages in the diagenesis of algal material in lakes, microbial attack was allowed to proceed on natural populations of two blue-green algal species, Gloeotrichia echinulata (J. E. Smith) P. Richter and Oscillatoria agardhii var. isothrix Skuja. Changes in composition and abundance of the major lipid fractions were related to differences in the ease of microbial attack and to the effect of oxygen on the decay process. The loss of labile unsaturated compounds during diagenesis is consistent with the small amount of these compounds in most lake sediments.     

Environmental factors influencing chlorophyll v. nutrient relationships in lakes

SUMMARY. 1. A model relating log chlorophyll a concentration to log epilimnetic total phosphorus (TP) concentration was re-examined based on: (a) comparative and temporal studies of four stratifying Wisconsin and other highly eutrophic temperate lakes; (b) comparative summer lake surveys from Iowa and Alberta.
2. Although P-limited, deeper lakes with long hydraulic residence times and low external and internal nutrient loading in summer had summer chlorophyll a yields below model predictions based on spring and summer epilimnetic TP concentrations.
3. For lakes with summer epilimnetic TP between 30 and 80 mg m⁻³, chlorophyll a concentrations exceeded model predictions based on summer TP. This relationship held even for Lake Delavan, Wisconsin, where the ratio of available N to P was unfavourably low during spring turnover, and where the trans-thermocline N:P flux ratio was sub-optimal for algal needs in early summer.
4. With increasing summer TP concentrations and/or increasing epilimnetic circulation depth (>5m), chlorophyll a concentrations fell below model predictions—independent of the potential for N-limitation. This plateauing in chlorophyll a response occurred at lower epilimnetic TP content (−2) in lakes with elevated non-algal light extinction coefficients. Using Tailing's algorithm for the'column compensation point' (algal photosynthesis = algal respiration over diel cycle), light limitation best explains this fall-off in chlorophyll a yield.
5. The failure of the Dillon & Rigler (1974) spring TP v . summer chlorophyll a model for these Wisconsin lakes is unrelated to N-limitation. Instead, it reflects internal adjustment in take TP in response to stratification and seasonal external P loading.     

Seasonal pattern in nutrient limitation and grazing control of the phytoplankton community in a non-stratified lake

1. The relative importance of zooplankton grazing and nutrient limitation in regulating the phytoplankton community in the non-stratified Lake Kvie, Denmark, were measured nine times during the growing season.
2. Natural phytoplankton assemblage bioassays showed increasing importance of nutrient limitation during summer. Growth rates at ambient nutrient concentrations were continually below 0.12 per day, while co-enrichment with nitrogen (N) and phosphorus (P) to above concentration-saturated conditions enhanced growth rates from May to the end of July.
3. Stoichiometric ratios of important elements in seston (C : N, C : P, N : P), in lake water (TN : TP), in external loading (TN : TP) and in internal loading (DIN : DIP) were measured to determine whether N or P could be the limiting nutrient. TN : TP molar ratio of both lake water, benthic fluxes and external loading suggested P limitation throughout the growing season. However, seston molar ratios suggested moderate P-deficiency only during mid-summer.
4. Abundance and community structure of the zooplankton varied considerably through the season and proved to be important in determining the responses of algal assemblages to grazing. High abundance of cladocerans and rotifers resulted in significant grazing impact, while cyclopoid copepods had no significant effect on the phytoplankton biomass.
5. Regeneration of ammonium and phosphate by zooplankton were periodically important for phytoplankton growth. A comparison of nutrient regeneration by zooplankton with nutrient inputs from sediment and external sources indicated that zooplankton may contribute significantly in supplying N and P for the growth of phytoplankton.     

The past and future of phytoplankton in the UK's largest lake,Lough Neagh

Lough Neagh is the largest lake in the UK and has been extensively monitored since 1974. It has suffered from considerable eutrophication and toxic algal blooms. The lake continues to endure many of the symptoms of nutrient enrichment despite improvements in nutrient management throughout the catchment, in particular a permanently dominant crop of the cyanobacterium Planktothrix agardhii. This study examines the historical changes in the Lough, and uses the PROTECH lake model to predict how the phytoplankton community may adapt in response to potential future changes in air temperature and nutrient load. PROTECH was calibrated against 2008 observations, with a restriction on the maximum simulated mixed depth to reflect the shallow nature of the lake and the addition of sediment released phosphorus throughout the mixed water column between 1 May and 1 October (with an equivalent in-lake concentration of 2.0 mg m⁻³). The historical analysis showed that phytoplankton biomass (total chlorophyll a) experienced a steady decline since the mid-1990s. During the same period the key nutrients for phytoplankton growth in the lake have shown contrasting trends, with increases in phosphorus concentrations and declines in nitrate concentrations. The modelled future scenarios which simulated a temperature increase of up to 3 °C showed a continuation of those trends, i.e. total chlorophyll a and nitrate concentrations declined in the surface water, while phosphorus concentrations increased and P. agardhii dominated. However, scenarios which simulated a 4 °C increase in air temperature showed a switch in dominance to the cyanobacteria, Dolichospermum spp. (formerly Anabaena spp.). This change was caused by a temperature related increase in growth driving nutrient consumption to a point where nitrate was limiting, allowing the nitrogen-fixing Dolichospermum spp. to gain sufficient advantage. These results suggest that in the long term, one nuisance cyanobacteria bloom may only be replaced by another unless the in-lake phosphorus concentration can be greatly reduced.     

Changes in phytoplankton physiology and morphology in response to dissolved nutrients in Lough Neagh, N. Ireland

SUMMARY. Cell phosphorus and anthrone-reactive carbohydrate contents were measured over a 6-month period in the phytoplankton of Lough Neagh, N. Ireland. When all the soluble reactive phosphorus had disappeared from the water column, cell phosphorus contents began to decrease, reaching a minimum value of 0.4% ash-free dry weight. At this time, cell carbohydrate contents greatly increased and phosphorus sorption assays showed rapid uptake of orthophosphate. Following a presumed sediment phosphorus release, cell carbohydrate and phosphorus uptake rate decreased greatly and there was a subsequent increase in cell phosphorus content to 1.6% ash-free dry weight. As the cell phosphorus content declined, blue-green algal filaments became progressively shorter. On the basis of these responses it is likely that phosphorus limitation occurred over a 4-week period.     

The relationship between nutrients and trophic‐level biomass in turbid tropical ponds

1. The total phosphorus–algal biomass relationship from a set of turbid tropical ponds in Kenya was compared with predictions derived from surveys of temperate and subtropical lakes. Despite high concentrations of total phosphorus (TP) (up to 797 μg L ^–1) and inorganic turbidity (up to 800 mg L^–1), the log–log relationship between algal biomass and TP was steeper than expected.
2. No evidence of nitrogen limitation was found at high TP, and total nitrogen (TN):TP ratios were higher than in lakes with similar TP levels studied previously. High TN:TP ratios may be a consequence of excretion by cattle into the ponds, a nutrient source characterized by a high ratio of available N to available P.
3. Despite extremely high turbidity, the ratio of mixed layer depth to euphotic depth was generally low because these ponds are shallow (≤ 2 m), and was not related to algal yield. A positive relationship was also found between TP and zooplankton biomass, and between TP and the density of the zooplanktivorous bug, Anisops . In contrast, no relationship was found between fish biomass and TP, algal biomass or zooplankton biomass.     

SEASONAL CYCLING OF ALGAL NUTRIENT LIMITATION IN CHAUTAUQUA LAKE,NEW YORK1

Algal nutrient enrichment bioassays were conducted between May 1975 and August 1978 using water samples collected from Chautauqua Lake, New York. Photosynthetic fixation rates of natural phytoplankton assemblages were enhanced by additions of phosphorus and nitrogen, although enrichment with other nutrients had no significant stimulatory effect on algal photosynthesis. Whereas phosphorus stimulated in spring and early summer, both nitrogen and phosphorus enhanced photosynthesis in midsummer and fall. Relative to the effect of phosphorus enrichment, enhancement of photosynthesis by nitrogen during the summer and fall was highest in the northern part of the lake. During the period of ice cover, photosynthesis did not appear to be limited by nutrients in that nutrient additions (P, N, Si, C, Fe, trace metals) did not enhance fixation rates. Observed temporal fluctuations in the response of the algae to P and N correlated with changes in the lake water N:P ratio as well as with temporal changes in dissolved orthophosphate and nitrate-nitrite nitrogen. The N:P ratio decreased drastically in the summer and remained at ca. 10 or less through mid-fall, suggesting that N concentrations were inadequate for the non-N-fixing phytoplankton. Studies over 3 yr indicate that states of P and N limitation undergo time-space fluctuations that occur in a cyclic pattern in the surface waters of Chautauqua Lake.     

Phosphorus and nitrogen limitation of phytoplankton growth in eutrophic Lake Inba, Japan

Lake Inba is one of the most eutrophic lakes in Japan. In this study, field sampling and nutrient enrichment bioassays were conducted to determine the seasonal patterns of nutrient limitation for phytoplankton growth in this lake. Phytoplankton biomass increased significantly with the additions of phosphorus (P) on almost all sampling dates, indicating P limitation of phytoplankton growth from spring to autumn. However, nitrogen (N) limitation was also observed during summer (i.e., 19 August). On 10 August, a typhoon struck Lake Inba. After this event, dissolved inorganic nitrogen (DIN) and phosphorus concentrations increased, probably because of increased river discharge. At the same time, phytoplankton growth in the control treatment became relatively high, with the addition of neither P nor N stimulating the growth. However, 10 days after the typhoon, the phytoplankton growth rate in the control treatment decreased, with only the addition of N having a significant positive effect on phytoplankton growth. N limitation during summer is caused by the low concentrations of DIN, as well as changes in the N:P ratio due to allochthonous nutrient loads. These results indicate that a reduction of both P and N input is necessary to control phytoplankton blooms in Lake Inba.     

珠江口及毗邻海域营养盐对浮游植物生长的影响

基于2006年7月(夏季),10月(秋季)和2007年3月(春季)的现场调查数据,对珠江口及毗邻海域中的营养盐和叶绿素a等环境生态因子的时空分布特性进行了对比分析,研究了氮磷比与叶绿素a含量和种群多样性之间的联系,探讨了该海域营养盐对于浮游植物生长的影响。结果表明:(1)研究海域营养盐表现出较强的季节和空间差异性,总氮(TN)和总磷(TP)浓度均值春季(1.545 mg/L、0.056 mg/L)和夏季(1.570 mg/L、0.058 mg/L)均大于秋季(1.442 mg/L、0.034 mg/L),且春夏季浓度空间差异更明显。(2)调查期间海域营养盐含量超标现象突出,夏季尤为明显。无机氮(DIN)总体均值0.99 mg/L,超四类海水标准限值1倍,活性磷酸盐(PO4-P)总体均值0.021 mg/L,DIN∶PO4-P平均值为130;叶绿素a浓度与营养盐、p H、温度有较显著的相关性。(3)叶绿素a浓度较高的站位,具有较高的DIN∶PO4-P值,但浮游植物多样性指数偏低,优势种明显,主要为中肋骨条藻。氮磷比的改变会影响不同生长特性的浮游植物间的竞争和种群结构的改变;今后海洋污染治理中,在控制氮、磷污染时要注意氮磷比的改变可能造成的浮游生态影响。     

Meteorological influences on algal bloom potential in a nutrient-rich blackwater river

1. The effect of variability in rainfall on the potential for algal blooms was examined for the St Johns River in northeast Florida. Water chemistry and phytoplankton data were collected at selected sites monthly from 1993 through 2003. Information on rainfall and estimates of water turnover rates were used in the analyses of trends in phytoplankton biomass.
2. Major trends in rainfall and runoff within the lower St Johns River catchment over the 10-year study period were marked by both significant drought and flood periods. Autumn and winter rainfall patterns were strongly correlated with the range of Pacific sea surface temperature anomalies associated with El Niño events and La Niña periods. The effect of these major shifts in rainfall was evident in the strong relationship to replacement rates for water within the lower St Johns River.
3. The eutrophic status of the river was reflected in the high concentrations of nitrogen and phosphorus observed at all sampling sites, with total nitrogen concentrations up to 3100  μ g L⁻¹ and total phosphorus concentrations up to 180  μ g L⁻¹.
4. While it is clear that the high phytoplankton biomass and frequent blooms that characterize the freshwater portions of the lower St Johns River are fundamentally based on nutrient status, the expression of that potential was strongly correlated to water replacement rates, as revealed by the inverse relationship between phytoplankton biovolume increase and water turnover rate, with an R ² of 0.80 for the major bloom season. The sensitivity of algal blooms to rainfall patterns over the 10-year study period suggest that longer-term temporal and spatial shifts in rainfall, such as multi-decadal cycles and the global-warming phenomenon, will also influence the frequency and intensity of algal blooms.     

Whitemere, a lake that defies some conventions about nutrients

1. Whitemere, a kettle-hole lake in north-west England (z_m,14 m, area 22.5 ha) has extremely high maximal concentrations (around 1 mg L⁻¹) of total phosphorus (TP) and soluble reactive phosphorus (SRP), and comparatively low maximal concentrations of dissolved inorganic nitrogen (<0.5 mg N L⁻¹). Bioassays indicate that its phytoplankton is nitrogen limited, and it has surface blue-green algal blooms in summer. Palaeoecological investigations have shown that high populations of cyanophytes occurred more than 6000 years ago, even before human settlement of the catchment.
2. The reasons for this combination of features are not immediately clear. This paper attempts to distinguish between two hypotheses: that the high phosphorus concentrations arise from input of phosphate-rich groundwater; and that the high phosphorus concentrations arise from concentration and recycling mechanisms within the lake.
3. The lake is entirely fed by ground water, direct rainfall and surface sheet flow. The concentrations of phosphorus in the ground water are much lower than in the lake water. The lake is stratified and has high phosphorus concentrations in the anaerobic hypolimnion in summer as a result of substantial release from the sediment. Vertical turbulence in the relatively weakly stratified water column may transfer substantial amounts to the epilimnion.There is also substantial phosphorus release from the sediments under the aerobic epilimnion.
4. There is little outflow of water and phosphorus. Most of the phosphorus is returned to the sediment under winter isothermal conditions, to be released again the following summer. The lake has probably maintained these mechanisms for millennia. Recent eutrophication is likely to have been driven by nitrogen inputs from greater intensification of agriculture in the catchment.     

Erosion,phosphorus and phytoplankton response in rivers of South-Eastern Norway

The development of P fractions and phytoplankton was studied in three rivers with varying concentrations of seston.Less than 1% of the yearly TP transport may take place during periods with high algal biomass.The observation of a high growth rate of phytoplankton in the rivers coinciding with high concentrations of RP, low content of seston and high TP:Chl a ratio, indicate that the growth was often not P-limiting. During short periods with high phytoplankton biomass the ratio TP:Chl a may be low, indicating that a high fraction of TP was available.The content of P in soil samples and in samples with high seston content was about 0.1% of dry weight, and the algal availability of P often varied between 25 and 75% of TP for both types of samples.Decreasing biomass or low growth rates were observed at secchi depths less than 0.5 m and seston concentrations less than about 25 mg dry weight 1^–1. High flow rate also depressed the development of the total phytoplankton biomass. The assimilation of available P is incomplete under such conditions, i.e. under conditions of light limitation and high dilution rate.The availability of P for phytoplankton in rivers with different length, light conditions and stream velocity is discussed.     

Temporal and spatial variations of chemical oxygen demand in Lake Taihu,China, from 2005 to 2009

We undertook a study in Lake Taihu, China, from 2005 to 2009 including a total of 639 samples to determine: (i) the seasonal dynamics and spatial distribution of the chemical oxygen demand (COD) and (ii) the relationships between the COD concentration and the biochemical oxygen demand (BOD), phytoplankton pigment, total dissolved nitrogen (TDN), and total dissolved phosphorus (TDP) concentrations, as well as the chromophoric dissolved organic matter (CDOM) absorption coefficient. There were significant spatial differences in the COD concentration, which gradually decreased from Zhushan Bay in the northwest, to the north, the lake center, and the southeast of the lake. The COD concentration was significantly higher at near-shore sites than that at open water sites. The mean COD concentrations were significantly higher in the spring and summer than in the winter and autumn. The lowest annual mean COD concentration appeared in 2009, which could be attributed to improvements in water quality management and high rainfall. The COD concentrations in all four seasons were strongly correlated with phytoplankton pigment, suggesting that extracellular release of COD from phytoplankton was an important COD source. The correlation coefficients between the COD and phytoplankton pigment concentrations were higher in the spring, summer, and autumn than in the winter, showing a more important contribution of phytoplankton degradation to COD in the algal bloom season than in the non-algal bloom season. These new data on the temporal and spatial characteristics of the COD in Lake Taihu will be crucial for developing future strategies for water quality management.     

A Long Term Study of the Phosphorus Cycle in Lough Neagh,Northern Ireland

Eleven years of weekly observations on the phosphorus cycle in Lough Neagh are analysed. The data comprised inputs, outputs and lake concentrations of the soluble and particulate phosphorus fractions. It is shown that the particulate phosphorus input quickly sediments to the bottom and the particulate phosphorus in the lake is largely made up of phytoplankton. The average phosphorus retention in the lake was 34 %, similar to the particulate phosphorus input. Climate has a big influence on the phosphorus input and sediment release, both of which varied two-fold. A simulation model suggested that suppression of sediment P release would reduce lake winter phosphorus concentrations by an average of 29 %.     

Essential fatty acid content and the phosphorus to carbon ratio in cultured algae as indicators of food quality for Daphnia

1. A series of experiments examined growth of Daphnia magna on three algal diets ( Rhodomonas minuta , Scenedesmus acutus and Synechococcus sp.) at varying physiological states [nitrogen and phosphorus (P) limitations] to test whether variation in algal fatty acid and/or elemental composition can predict Daphnia growth.
2. These algae differed widely in their essential fatty acid (EFA) composition while phosphorus (P) or nitrogen limitation had only a small influence on their ω3-polyunsaturated fatty acid (PUFA) content.
3. Individually, algal ω3-PUFA content explained 69% of the variation in the normalised growth of D. magna, while algal phosphorus content explained 11% of the variation. Quantitative models for D. magna growth used both algal ω3-PUFA content and algal C : P ratio as food quality indices. Together, algal ω3-PUFA content and C : P ratio explained 70% of the variation in the normalised growth rate of D. magna .
4. Our results indicate that EFA influenced algal food quality much more strongly than P. The EFA and mineral P impacts appear to be independent.     

Phytoplankton, light and nutrients along a gradient of mixing depth: a field test of producer-resource theory

1. Variation in depth of the mixed surface layer of temperate lakes should affect phytoplankton dynamics because, with increasing mixing depth, average light intensity in and specific sedimentation losses out of the mixed layer both decrease. 2. Our aim was to test a recent dynamic model which relates phytoplankton biomass and the availability of production‐limiting resources (light and dissolved mineral nutrients) to mixing depth and nutrient supply from external sources. 3. During summer stratification we sampled the mixed layers of 30 dimictic, phosphorus‐limited, oligo‐ to mesotrophic, mostly non‐humic lakes north of the Alps. 4. The results agree well qualitatively with model expectations. Algal concentration in the mixed layer was negatively related to mixing depth or its surrogate log‐transformed lake area. Light intensity at the bottom of the mixed layer decreased whereas the concentration of available, inorganic phosphorus increased with increasing mixing depth. Across all depths, higher total phosphorus content was accompanied by higher phytoplankton biomass, lower light availability, and higher inorganic phosphorus concentration. 5. Our data match the predicted shift with increasing mixing depth from predominantly nutrient limitation towards increased light limitation of algal biomass.     

Can a community of small-bodied grazers control phytoplankton in rivers?

1. Phytoplankton, zooplankton and grazing were monitored throughout the growing season for three years (1994–96) in the Belgian section of the River Meuse.
2. A size structure analysis of the algal community shows that there was a summer shift toward larger algal units, following a decline in phytoplankton biomass. These changes occurred after an increase in zooplankton biomass and diversity.
3. Daily filtration rates of grazers ranged from 1 to 113% day^–1 and maxima were observed during the summer period. Higher rates tended to correspond with peaks of rotifer biomass. A decline in total phytoplankton biomass within two weeks followed the increase in zooplankton biomass and filtration rate. A rapid biomass recovery was then observed, along with a shift of the algal community toward larger units. When grazing activity was not sustained, due to zooplankton fluctuations, the change in phytoplankton size structure was less marked.
4. We suggest that the composition of the phytoplankton community of large rivers may at times be controlled by grazers. However, such biotic interactions can take place only when physical constraints are reduced, i.e. when discharge is low, and when increased transfer time, high temperature and availability of grazeable algae allow high zooplankton biomass.     

Variation in nutrient limitation of lotic and lentic algal communities in a Texas (USA) river

Nutrient limitation of periphyton and phytoplankton was assessed in the Upper Guadalupe River, Texas USA. Nutrient-diffusing substrates with added nitrogen (N) and phosphorus (P) were used to identify the limiting nutrient for lotic algae at three river sites in summer, fall, and winter. Pots enriched with P had significantly higher chlorophyll a concentrations for 7 of 9 trials. Added N alone did not significantly increase algal standing crops, although it was found to be secondarily limiting on one (and possibly two) occasions. Flow-through enrichment experiments were conducted in order to quantify the concentration of P needed to significantly increase algal standing crops. Response to enrichment was rapid when ambient P concentration was low (< 0.010 mg L^–1), but more moderate when ambient P levels were higher (0.015–0.025 mg L^–1). Nutrient limitation of phytoplankton in small surface-release reservoirs varied throughout the study, but N was either primarily or secondarily limiting in 6 of 8 trials; shifts in the limiting nutrient were correlated with fluctuations in flow into the reservoirs. Our enrichment studies show that algal response to nutrient addition was unpredictable as phytoplankton tended to be N-limited while periphyton was mainly P-limited. Further, while discharge apparently dictated the nutrient-biomass relationship for phytoplankton in reservoirs, ambient nutrient level is an important determinant of lotic periphyton response to enrichment.     

Lake‐type‐specific seasonal patterns of nutrient limitation in German lakes,with target nitrogen and phosphorus concentrations for good ecological status

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