The phytoplankton of Great Bitter Lake,Egypt, including the impacts of nutrient-laden and heated effluents

A year-long survey of the phytoplankton and nutrients in Great Bitter Lake indicates that this is a severely eutrophic lake. Chlorophyll levels were consistently high (>30μg/l), especially in summer (>90μg/l). The phytoplankton community comprised mostly diatoms and blue-green algae, although dinoflagellates and green algae were important at times. Local effects of effluent from a drain coming from the city of Ismailia were evident, although the effect of tourist hotels at Palma Beach was not detectable. The discharge of heated water from a thermal power plant raised the water temperature in the impacted area by 13°C in autumn and 22°C in summer and also caused decreases in chlorophyll and phytoplankton abundance. Dissolved nutrient levels were high and, especially at the offshore station, had a rather constant ratio of dissolved inorganic nitrogen to phosphate. The distribution of phytoplankton and chlorophyll were related more to temporal, presumably seasonal, variation than proximity to point sources of pollution.     

余氯对水生生物的影响

氯是滨河、滨海企业冷却水常用的防治污损生物的处理剂。总结近年来国内外氯在企业冷却水中的应用、氯对浮游植物、浮游动物、贝类、鱼类等影响的研究成果,为制定冷却水余氯排放标准和水产养殖的合理布局提供了参考,针对我国大量滨海电厂即将建立的现状,分析了余氯研究在中国海域研究的不足,提出了氯对我国海洋生物影响不同层面的研究方向。认为氯对生物种群的毒性,氯对生物群落组成、结构和生态演替的影响,以及减轻或避免余氯污染的对策是需要进一步解决的科学问题。     

热冲击和加氯后亚热带海区浮游植物细胞数量的动态变化

 近年来, 随着电厂在河口和港湾等生态脆弱区和敏感区的急剧增多, 电厂冷却系统热冲击和加氯对浮游生物造成的伤害已成为沿海地区非常严峻的生态安全问题。为探明冷却系统升温和加氯联合作用对亚热带海区浮游植物的影响程度, 针对滨海电厂的实际运作情况, 在室内对采自浙江省乐清湾海域的四季浮游植物进行了不同水平的热冲击和加氯胁迫, 并观察了这些浮游植物细胞数量在15 d内的动态变化。结果表明, 热冲击、加氯和季节均显著影响浮游植物细胞数量的恢复(p＜0.001), 其中, 加氯的影响最大, 季节次之, 热冲击影响最小, 但热冲击增强了氯对浮游植物的毒性。自然水温越高、升温幅度越大, 细胞数量恢复越慢。春、秋、冬季自然水温较低时, 升温4～12 ℃后, 细胞数量仅需1～6 d即可恢复到对照组水平; 夏季自然水温较高, 升温4～8 ℃后, 细胞数量需4～9 d恢复到对照组水平, 但升温12 ℃后, 细胞数量在15 d内未能恢复到对照组水平。加氯浓度越高, 细胞数量恢复越慢。加氯1.0～1.8 mg•L^−1后, 浮游植物生长虽受影响, 但大多能在15 d内恢复; 而加氯5.6 mg•L^−1后, 其生长受到完全抑制, 细胞数量在15 d内未恢复到对照组水平。     

Effects of Nutrients,Temperature and Their Interactions on Spring Phytoplankton Community Succession in Lake Taihu,China

We examined the potential effects of environmental variables, and their interaction, on phytoplankton community succession in spring using long-term data from 1992 to 2012 in Lake Taihu, China. Laboratory experiments were additionally performed to test the sensitivity of the phytoplankton community to nutrient concentrations and temperature. A phytoplankton community structure analysis from 1992 to 2012 showed that Cryptomonas (Cryptophyta) was the dominant genus in spring during the early 1990s. Dominance then shifted to Ulothrix (Chlorophyta) in 1996 and 1997. However, Cryptomonas again dominated in 1999, 2000, and 2002, with Ulothrix regaining dominance from 2003 to 2006. The bloom-forming cyanobacterial genus Microcystis dominated in 1995, 2001 and 2007–2012. The results of ordinations indicated that the nutrient concentration (as indicated by the trophic state index) was the most important factor affecting phytoplankton community succession during the past two decades. In the laboratory experiments, shifts in dominance among phytoplankton taxa occurred in all nutrient addition treatments. Results of both long term monitoring and experiment indicated that nutrients exert a stronger control than water temperature on phytoplankton communities during spring. Interactive effect of nutrients and water temperature was the next principal factor. Overall, phytoplankton community composition was mediated by nutrients concentrations, but this effect was strongly enhanced by elevated water temperatures.     

象山港电厂温排水增温对浮游细菌群落空间分布的影响

沿海电厂产生的温排水已造成了较严重的环境问题,如生物病害和赤潮暴发频率增加。现有研究多关注温排水增温对浮游动、植物多样性的影响,而缺乏在物质能量循环过程起核心纽带作用的微生物对增温的响应和反馈。考虑到生态系统对增温的反馈取决于浮游植物的初级生产力和微生物异养呼吸之间的平衡,利用Illumina测序技术结合水体理化性质研究了象山港电厂温排水增温梯度下浮游细菌的空间分布特征。温排水预期的海水增温显著地增加了水体中硝氮(P=0.041,单因素方差分析)、化学需氧量(P0.001)、油污(P=0.004)和余氯(P=0.003)的浓度;但降低了溶解氧(P=0.034)和叶绿素a(P=0.045)的含量。此外,相似性分析发现温排水增温显著地(r=0.338;P=0.042)改变了浮游细菌群落结构,空间分布遵循空间距离-群落相似性衰减(r=-0.582;P=0.026)模型,周转速率为0.0013。细菌多样性主要受水体溶解氧、化学需要量和叶绿素a的影响,分别控制了34.6%、20.1%和10.0%的多样性变异。冗余分析(RDA)群落变异主要受环境因子(包括温度、油污、溶解氧和叶绿素a)的影响,一共解释了55.6%的群落变异;增温仅解释了4.8%的群落变异。因此,浮游细菌群落结构主要受环境因子的影响,这种各因子对群落变异相对贡献比例的特征与增温主要通过改变水质和浮游植物特征,以间接作用影响细菌群落组成的观念一致。此外,细菌多样性和群落组成受不同环境因子的驱动。相比而言,空间距离只控制了较小比例的群落变异(7.1%)。此外,筛选到11个细菌科,这些科的相对丰度与增温幅度显著相关,变化特征与各科已知的生态功能相吻合,如海洋螺菌科(Oceanospirillaceae)中有些菌株能够降解石油污染物,其相对丰度在高油污浓度站点增加(油污浓度与增温幅度正相关,r=0.558;P=0.030);嗜温的弧菌科相对丰度与增温幅度正相关。综上,研究结果初步明确了浮游细菌群落对电厂温排水增温的响应特征,并筛选到敏感的细菌科来指示和预测增温对生态功能的潜在影响。     

Statistical analysis of phytoplankton biomass in coastal waters: Case study of the Wadden Sea near Lauwersoog (The Netherlands) from 2000 to 2009

As the most important indicator of the coastal ecosystem, various studies on phytoplankton have frequently been proposed in coastal waters. In this study, the statistical analysis of phytoplankton biomass (in terms of chlorophyll a) is performed in the Wadden Sea near Lauwersoog (53.4167° N, 6.2000° E), The Netherlands, determined by the 10-year's historical dataset from 2000 through 2009. Boxplot analysis is introduced to give insight in the seasonal variations of phytoplankton biomass and physical–chemical conditions. Annually, a big difference is found in chlorophyll a, showing that the maximum values occur in the months of April and July, and the minimum values occur in winter (December, January, and February). The phytoplankton biomass is positively correlated with the physical conditions (salinity, light intensity, and water temperature), and is negatively correlated with the nutrients (nitrate, ammonium, and silicate). Factor analysis distinguishes the driving variables that represent most of the total variance using three extraction methods (principal component analysis, unweighted least squares, and maximum likelihood). Nitrate dominates the main activity in the first rotated component/factor, and ammonium contributes the most in the second rotated component/factor. The findings of this study are significant for understanding the roles of the environmental conditions upon the phytoplankton variability, and for reducing the overlapping information to characterize the driving variables.     

滨海电厂冷却水余热和余氯对中华哲水蚤的影响

依据滨海电厂冷却系统的实际运作情况,模拟研究了滨海电厂冷却水余热和余氯对中华哲水蚤的影响.结果表明：中华哲水蚤的热忍受能力随暴露时间延长而降低,随驯化温度升高而升高.驯化温度为16 ℃～27 ℃的中华哲水蚤在持续受到15、30、45 min热冲击及持续升温暴露24和48 h的致死温度分别为29.9 ℃～31.7 ℃、29.4 ℃～31.0 ℃、28.9 ℃～30.3 ℃和26.9 ℃～28.5 ℃、26.4 ℃～28.0 ℃.当驯化温度升高到一定程度后,其热忍受能力不再上升.电厂冷却水中余氯对中华哲水蚤的毒性随驯化温度升高、升温幅度增大及暴露时间延长而增强.     

Filamentous green algae inhibit phytoplankton with enhanced effects when lakes get warmer

1. Filamentous green algae (FGA) may represent an alternative state in high‐nutrient shallow temperate lakes. Furthermore, a clear water state is sometimes associated with the dominance of FGA; however, the mechanisms involved remain uncertain. 2. We hypothesised that FGA may promote a clear water state by directly suppressing phytoplankton growth, mostly via the release of allelochemicals, and that this interaction may be affected by temperature. 3. We examined the relationships between FGA, phytoplanktonic chlorophyll a concentrations and zooplankton in a series of mesocosms (2.8 m³) mimicking enriched shallow ponds now and in a future warmer climate (0 and c. 5 °C above ambient temperatures). We then tested the potential allelopathic effects of FGA (Cladophora sp. and Spirogyra sp.) on phytoplankton using several short‐term microcosms and laboratory experiments. 4. Mesocosms with FGA evidenced lower phytoplanktonic chlorophyll a concentrations than those without. Zooplankton and zooplankton : phytoplankton biomass ratios did not differ between mesocosms with and without FGA, suggesting that grazing was not responsible for the negative effects on phytoplanktonic biomass (chlorophyll a). 5. Our field microcosm experiments demonstrated that FGA strongly suppressed the growth of natural phytoplankton at non‐limiting nutrient conditions and regardless of phytoplankton initial concentrations or micronutrients addition. Furthermore, we found that the negative effect of FGA on phytoplankton growth increased up to 49% under high incubation temperatures. The experiment performed using FGA filtrates confirmed that the inhibitory effect of FGA on phytoplankton may be attributed to allelochemicals. 6. Our results suggest that FGA control of phytoplankton growth may be an important mechanism for stabilising clear water in shallow temperate lakes dominated by FGA and that FGA may play a larger role when lakes get warmer.     

The response of phytoplankton community to anthropogenic pressure gradient in the coastal waters of the eastern Adriatic Sea

In order to test the response of phytoplankton to anthropogenic pressure, data of chlorophyll a concentration, phytoplankton abundance, and composition are analyzed in relation to anthropogenic pressure gradient and environmental variables such as temperature, salinity and nutrients. Investigated sites encompassed wide tropic range according to a preliminary determination of anthropogenic pressure, quantified through the LUSI index. Statistical analyses indicated nitrates and silicates as proxies of freshwater influence, and phytoplankton single metrics such as concentrations of chlorophyll a and abundances as indicators of anthropogenic pressure. Boundary values for different water quality classes for coastal waters under indirect freshwater influence (Type II) are obtained according to gradient between concentration of chlorophyll a and pressure index (LUSI), which empirically fit to exponential equation. The response of phytoplankton diversity was not linear, as the highest diversity was observed in the area with intermediate disturbance level. CCA analysis identified Skeletonema marinoii, Scrippsiella trochoidea, Guinardia flaccida, Leptocylindrus spp., Prorocentrum spp., Proboscia alata, Eutreptiella spp., and Pseudonitzschia spp. as local eutrophication indicators, whose abundances increased with nutrients loads.     

Temperature-Correlated Changes in Phytoplankton Community Structure Are Restricted to Polar Waters

Globally distributed observations of size-fractionated chlorophyll a and temperature were used to incorporate temperature dependence into an existing semi-empirical model of phytoplankton community size structure. The additional temperature-dependent term significantly increased the model’s ability to both reproduce and predict observations of chlorophyll a size-fractionation at temperatures below 2°C. The most notable improvements were in the smallest (picoplankton) size-class, for which overall model fit was more than doubled, and predictive skill was increased by approximately 40%. The model was subsequently applied to generate global maps for three phytoplankton size classes, on the basis of satellite-derived estimates of surface chlorophyll a and sea surface temperature. Polar waters were associated with marked decline in the chlorophyll a biomass of the smallest cells, relative to lower latitude waters of equivalent total chlorophyll a. In the same regions a complementary increase was seen in the chlorophyll a biomass of larger size classes. These findings suggest that a warming and stratifying ocean will see a poleward expansion of the habitat range of the smallest phytoplankton, with the possible displacement of some larger groups that currently dominate. There was no evidence of a strong temperature dependence in tropical or sub-tropical regions, suggesting that future direct temperature effects on community structure at lower latitudes may be small.     

Differential Sunlight Sensitivity of Picophytoplankton from Surface Mediterranean Coastal Waters

We tested the sensitivity of coastal picophytoplankton exposed to natural sunlight in short-term experiments. Cell abundance and cell-specific chlorophyll fluorescence were significantly reduced in Prochlorococcus spp. but not in Synechococcus, whereas picoeukaryotes had an intermediate response. These results are the first direct evidence of a differential sensitivity to sunlight of these ubiquitous marine members of unicellular phytoplankton.     

Phytoplankton growth under temperature stress: Laboratory studies using two diatoms from a tropical coastal power station site

(1) The paper reports results of laboratory studies on growth rates at elevated temperatures of two diatoms (Chaetoceros wighami and Amphora coffeaeformis) isolated from the intake and outfall sites of a coastal power station on the east coast of India. (2) Temperature exposures consisted of acute and chronic treatments to simulate condenser transit and prolonged thermal plume entrainment. (3) The results showed that the various temperature zones presented by cooling water circuit of the power station would permit diatom growth, with growth rates that are marginally different form those at average ambient sea temperature. (4) Fundamental differences in the response of different diatom species to temperature stress result in the abundance of the more tolerant species such as A. coffeaeformis which dominate in an elevated temperature zone, such as that offered by the discharge canal of the power station.     

Tolerance of copepods to short-term thermal stress caused by coastal power stations

The tolerance of marine copepods to short-term thermal stress was measured by the median lethal temperature (LT₅₀) tests in laboratory. Experiments on LT₅₀ of copepods from different acclimation and acclimatization conditions collected from the Yueqing Bay were carried out under heat exposure for 15, 30 and 45 min. The LT₅₀ of copepods decreased with increasing exposure time but increased with rising acclimation and acclimatization temperatures. However, the differences in copepod LT₅₀ decreased with rising acclimatization temperatures, which suggested that entrained copepod mortality increased with raised water temperature due to the acute thermal stress of coastal power stations. Results also revealed that the thermal tolerance of Labidocera euchaeta was much higher than that of Calanus sinicus in spring. The thermal tolerances of different copepod species in summer were in the order, Pseudodiaptomus marinus, Acartia spinicauda, Acartia pacifica and L. euchaeta.     

Grazing effect of the invasive reef-forming polychaete Ficopomatus enigmaticus (Fauvel) on phytoplankton biomass in a SW Atlantic coastal lagoon

In this work we evaluate the potential grazing impact of the invasive reef-forming polychaete Ficopomatus enigmaticus in a SW Atlantic coastal lagoon (Mar Chiquita coastal lagoon; 37° 40' S, 57° 23' W; Argentina). This gregarious species feeds on suspended detritus and phytoplankton. Given the large area dominated by reefs, suspension feeding by this species might reduce the lagoon phytoplankton concentration and even hinder local eutrophication. To evaluate this hypothesis in situ replicated mesocosm experiments were performed in spring 2005 and, summer and winter 2006. Mesocosms enclosing reefs and without reefs were installed and grazing intensity was measured as the difference in chlorophyll a concentration and turbidity between the reef-treatment and the treatment without reefs. Reefs of F. enigmaticus decreased the mean chlorophyll a concentration, more during summer (56% decrease) than spring (25% decrease) and winter (19% decrease). Reefs also decreased water turbidity during summer (54% decrease) and spring (23% decrease). While previous studies indicate that the physical structure of these reefs alters water flow increasing water turbidity, our evidences show that their suspension-feeding activity can counteract this effect. Water turbidity was positively correlated with chlorophyll a concentration, which suggests that phytoplankton grazing by F. enigmaticus decreased light attenuation through the water column, with a potential for enhancement of benthic primary productivity. Therefore, our results suggest that grazing by this polychaete affects overall estuarine primary production as well as the relative importance of planktonic and benthic carbon sources to higher trophic levels.     

The synergism between hydrocarbon pollutants and UV radiation: a potential link between coastal pollution and larval mortality

Coastal, benthic invertebrates with complex life history strategies are exposed to stage- and habitat-specific selective forces. In the coastal environment, benthic adults are exposed to polycyclic aromatic hydrocarbon pollutants (PAHs) due to their proximity to human activities (shipping, urbanization, and industrialization). Benthic invertebrates produce lipid-rich eggs or larvae that absorb PAHs from polluted estuaries and coastal waters. The larvae of many coastal invertebrates move offshore following release from benthic adults. During development in offshore waters, larvae of some species are exposed to relatively high levels of ultraviolet (UV) radiation. Marine organisms vary in their tolerance to PAHs and UV radiation. The purpose of this study was to examine the effects of the sequential exposure of the larvae of marine crabs to PAHs and UV radiation.Using laboratory experiments, the larvae of four crab species were exposed to PAHs and UV radiation. There was a significant synergistic effect of exposure to PAH (fluoranthene or pyrene) and UV radiation on larvae of the spider crab (Libinia dubia), the stone crab (Menippe adina) and the mud crab (Panopeus herbstii). Larvae of blue crabs (Callinectes sapidus) were exposed to PAHs and UV radiation in both laboratory and solar UV experiments. Significantly higher mortality occurred for C. sapidus larvae using either type of UV-artificial or solar.Larvae of coastal invertebrates with complex life history strategies are susceptible to the combined effects of PAHs and UV radiation. In this study, the exposure of crab larvae to PAHs and UV radiation resulted in mortality to crab larvae using laboratory and solar UV experiments. There were no effects on larval crab mortality due to PAH or UV radiation independently but mortality was as high as 100% when both factors were present.     

Complex seasonal patterns of primary producers at the land-sea interface

Seasonal fluctuations of plant biomass and photosynthesis are key features of the Earth system because they drive variability of atmospheric CO₂, water and nutrient cycling, and food supply to consumers. There is no inventory of phytoplankton seasonal cycles in nearshore coastal ecosystems where forcings from ocean, land and atmosphere intersect. We compiled time series of phytoplankton biomass (chlorophyll a) from 114 estuaries, lagoons, inland seas, bays and shallow coastal waters around the world, and searched for seasonal patterns as common timing and amplitude of monthly variability. The data revealed a broad continuum of seasonal patterns, with large variability across and within ecosystems. This contrasts with annual cycles of terrestrial and oceanic primary producers for which seasonal fluctuations are recurrent and synchronous over large geographic regions. This finding bears on two fundamental ecological questions: (1) how do estuarine and coastal consumers adapt to an irregular and unpredictable food supply, and (2) how can we extract signals of climate change from phytoplankton observations in coastal ecosystems where local‐scale processes can mask responses to changing climate?     

Application of a new multi-metric phytoplankton index to the assessment of ecological status in marine and transitional waters

Patterns of phytoplankton size spectra variation with gradients of environmental stress have been observed in freshwater, transitional waters and marine ecosystems, driving the development of size spectra based assessment tools.In this study, we have tested on transitional and coastal waters a new Index of Size spectra Sensitivity of Phytoplankton (ISS-Phyto), which integrates simple size spectra metrics, size class sensitivity to anthropogenic disturbance, phytoplankton biomass (chlorophyll a) and taxonomic richness thresholds. ISS-Phyto has been tested both among and within ecosystems along pressure gradients based on expert view assessment; the adequacy of symmetric and both left and right asymmetric models of phytoplankton size class sensitivity have been compared.The results showed that ISS-Phyto consistently discriminated between anthropogenic and natural disturbance conditions. Left asymmetric models of size spectra sensitivity, assuming greater disturbance tolerance with respect to eutrophication and organic enrichment of increasingly large size classes, showed the best fit comparing all ecosystems; in three of the four considered ecosystems (Varna, Helsinki, Mompás-Pasaia), they seemed to discriminate best between different levels of disturbance also within ecosystems. Moreover, they demonstrated significant and inverse patterns of variation along the overall pressure gradient as well as along the inorganic phosphorus (DIP), chlorophyll a and trophic index (TRIX) gradients.Therefore, ISS-Phyto, originally developed for transitional waters, seems to be an adequate assessment tool of ecological status also in coastal marine waters; moreover, it seems adequate to describe within ecosystem disturbance gradients. Hence, ISS-Phyto helps to understand the relationships between anthropogenic impact and ecosystem response from the individual point of view, with reference to the simple parameter of body size.     

Decadal weakening of the wind-induced upwelling reduces the impact of nutrient pollution in the Bay of Málaga (western Mediterranean Sea)

The hypothesis that nitrate versus phosphate regulates the coastal primary production has been assessed at different time scales in the northwest Alboran Sea. Time series of temperature, salinity, nutrients and chlorophyll a obtained at three stations located off Málaga city (the greatest coastal urban core along the Alboran Sea shoreline) from 1992 to 2006 have been analysed. At the decadal scale, temperature increased linearly while salinity decreased. These changes were related to a shift in the wind regime suggesting that coastal upwelling became steadily weaker. In contrast to phosphate, nitrate was positively correlated with salinity at the seasonal scale and decreased linearly from 1992 to 2006. Seasonal and decadal changes in chlorophyll a were correlated with nitrate (and uncorrelated with phosphate). However, non-regular variability in chlorophyll a was correlated with phosphate. Consequently, the results demonstrate that nitrate controls the phytoplankton biomass at the inter-annual scale while both nitrate and phosphate do so at a shorter time scale. The Bay of Málaga receives elevated entries of domestic waste waters that release high loads of phosphate compared to nitrate. Our analysis indicates that the expected impact of this pollution on chlorophyll a at the inter-annual scale is reduced in comparison with the effects of atmospheric forcing.     

Phytoplankton sterol contents vary with temperature,phosphorus and silicate supply: a study on three freshwater species

The understanding of environmentally induced changes in the biochemical composition of phytoplankton species is of great importance in both physiological studies and ecological food web research. In extensive laboratory experiments we tested the influence of two different temperatures (10°C and 25°C) and a phosphorus supply gradient on the sterol concentrations of the three freshwater phytoplankton species Scenedesmus quadricauda, Cryptomonas ovata and Cyclotella meneghiniana. The diatom C. meneghiniana was additionally exposed to a silicate gradient. In two separate experiments we analysed (1) possible interactive effects of temperature and phosphorus supply and (2) the effect of four phosphorus levels and three silicate levels on algal sterol concentrations. We observed that sterol concentrations were higher at 25°C than at 10°C in S. quadricauda and C. meneghiniana, but were not affected by temperature in C. ovata. Interactive effects of temperature and phosphorus supply on sterol concentrations were found in C. meneghiniana. This presumably was due to the bioconversion of one sterol (24-methylenecholesterol) into another (22-dihydrobrassicasterol). Increasing phosphorus supply resulted in species-specific effects on sterol concentrations, viz. an optimum curve response in S. quadricauda, a saturation curve response in C. meneghiniana and no change in sterol concentration in C. ovata. Effects of silicate supply on the sterols of C. meneghiniana equalled the effects of phosphorus supply. Albeit we did not observe a general trend in the three phytoplankton species tested, we conclude that sterol concentrations of phytoplankton are strongly affected by temperature and nutrient supply. Interactive effects point out the importance of taking into account more than just one environmental factor when assessing the effects of environmentally induced changes on phytoplankton sterol concentrations.     

Symbiotic association of the ciliate Ophrydium naumanni with Chlorella causing a deep chlorophyll a maximum in an oligotrophic South Andes lake

Vertical profiles of temperature, light and chlorophyll a concentration were examined in Lake Moreno Oeste, an oligotrophic South Andean lake (Argentina), during the warmest period of the year (November-April), when thermal stratification is characteristic. Concurrent samples for the enumeration of phytoplankton and green ciliates were taken, and the different contribution of these fractions to total chlorophyll a concentration was analysed. The development of a distinctive deep chlorophyll maximum was observed during summer months. The deep chlorophyll maximum was situated near the limit of the euphotic zone and just below the upper limit of the metalimnion. The results showed that the green ciliate Ophrydium naumanni with endosymbiotic Chlorella dominated the metalimnion causing the deep chlorophyll maximum. Additional laboratory experiments revealed a strong dependence of O.naumanni on light. Therefore, the symbiotic association appears to be an effective exploitation of the water column in poor-nutrient-high-light ecosystems like large Andean lakes.