灞河城市段浮游生物群落结构时空变化及其与环境因子的关系

为了解灞河城市段浮游生物群落结构及其与环境因子之间的关系,于2016年9月至2017年7月每2个月进行一次采样分析。调查期间共鉴定出浮游植物7门63属,其中绿藻门种类数最多(34.9%),其次为硅藻门(30.2%);浮游动物4类45种,以轮虫(48.9%)和原生动物(24.4%)为主。浮游植物和浮游动物的丰度分别为0.73×10~4—98.5×10~4个/L和20—1084个/L,在时空分布上均呈现下游高于上游的趋势,峰值均出现在夏季。根据水体透明度、溶解氧、总磷、总氮、高锰酸盐指数和重金属等理化指标、浮游生物丰度、Shannon-Wiener多样性指数和Margalef丰富度指数对灞河城市段水质状况进行综合评价,结果显示,所研究流域总体为中污状态,上游景观河道内的水生植物对水体具有一定的修复能力。典范对应分析表明,水温、总氮、pH、溶解氧和汞是浮游植物群落分布的主要影响因子;影响浮游动物群落分布的主要因子为水温、总氮、总磷、叶绿素a和汞。     

广东长潭水库浮游生物群落结构特征

长潭水库是广东梅州市重要的备用水源地,属于国家一类水源保护区。2010年7月,通过分析长潭水库浮游生物群落组成、生物量及多样性指数等群落结构特征和水体理化指标,揭示长潭水库富营养化水平。结果显示,长潭水库鉴定出55种浮游植物,隶属于6个门,优势种为绿藻门的栅藻（Scenedesmus sp.）、蓝藻门的微囊藻（Microcystis sp.）和伪鱼腥藻（Pseudanabaena sp.）、隐藻门的隐藻（Cryptophyta sp.）以及硅藻门的小环藻（Cyclotella sp.）和针杆藻（Synedra sp.）;藻类密度范围1.71×10⁶~2.44×10⁸ cells/L;浮游动物共检出44种,包括轮虫、枝角类和桡足类,其中轮虫是主要类群。综合营养状态指数评价结果显示,长潭水库部分水体呈轻度富营养化。典范对应分析结果显示,浮游动物、溶解氧、总氮、总磷和化学需氧量是影响长潭水库浮游植物群落结构的关键环境因子。     

Plankton composition and water quality in a pond of spring origin in Turkey

Phyto/zooplankton composition, chlorophyll a, and some water quality parameters were investigated in a spring-originated pond in Central Anatolia between February 2001 and January 2002. Water temperature, pH, dissolved oxygen, Secchi depth, total and calcium hardness, nitrate-nitrogen, nitrite-nitrogen, ammonia-nitrogen, total phosphorus, and soluble reactive phosphorus levels were analyzed. A total of 49 species belonging to Bacillariophyceae, Chlorophyceae, Cyanophyceae, Cryptophyceae, and Dinophyceae were identified. The highest phytoplankton abundance was found in August, whereas the lowest was determined in January. Phytoplankton abundance increased from February to August and declined in the following months. The Bacillariophyceae were dominant in the phytoplankton community. A total of 21 species of Rotifera, 2 species of Cladocera, and 1 genus of Copepoda were found. The zooplankton community was dominated by Rotifera. The highest abundance of zooplankton was recorded in July and the lowest value in November. The annual mean concentration of chlorophyll a was measured as 1.90 μg l⁻¹. In spite of these eutrophic levels (mean values of total phosphorus and nitrate-nitrogen: 0.069 mg P l⁻¹ and 0.68 mg N l⁻¹), phytoplankton cannot grow satisfactorily because of the short water retention time (0.6 day⁻¹). The shallowness of the pond together with the low phytoplankton biomass and the high concentrations of nutrients are discussed.     

Composition and variation of phytoplankton communities during Microcystis bloom in an artificial lagoon of Hangzhou Bay,China

The interaction of various environmental triggers on phytoplankton communities of an artificial lagoon of Hangzhou Bay China, was studied during a Microcystis bloom in summer 2016. Forty-two phytoplankton genera (six phyla) were defined, with Bacillariophyta accounting for half of all phytoplankton genera. It was determined that Melosira, Chlorella, Cyclotella, Microcystis, Merismopedia, Anabaena and Selenastrum, which were identified and counted by an inverted microscope, were the dominant genera. In addition, a series of environmental indicators were analyzed, including salinity, seawater temperature, dissolved inorganic nitrogen, soluble reactive phosphorus (PO₄-P), ammonium (NH₄-N), nitrate nitrogen (NO₃-N), nitrite (NO₂-N), silicate (SiO₄-Si), and chemical oxygen demand of the water samples, as well as zooplankton community. The results of variance partitioning by R language revealed that the most influential factor driving the change in the phytoplankton community was the environment (49.7%), and zooplankton grazing represented only 7.9%. The results of redundancy analysis indicated that the change and composition of the phytoplankton community correlated significantly with the interaction of salinity, PO₄-P, transparency, seawater temperature, and the dominant zooplankton species. Notably, salinity and temperature fluctuation were the key factors inducing the rapid succession of the plankton community in artificial lagoons such as within the Jinshan City Beach (Shanghai, China).

     

Temporal variations and pond age effect on plankton communities in semi-intensive freshwater marron (Cherax cainii,Austin and Ryan, 2002) earthen aquaculture ponds in Western Australia

The abundance and diversity of the plankton community represents the health of the aquatic ecosystem, and plays an important role in the growth of cultured animals under aquaculture conditions. The temporal variations of plankton abundance, taxonomic composition, diversity, evenness and species richness were studied in three old and three new semi-intensive marron (Cherax cainii, Austin and Ryan, 2002) ponds. Water parameters such as temperature, dissolved oxygen, pH, turbidity, TAN, nitrite, nitrate and reactive phosphate were recorded, and plankton samples were collected every two months, for one year of juvenile production cycle. A total of twenty-six phytoplankton and seven zooplankton genera were recorded. Chlorophyceae was the dominant class of phytoplankton throughout the year, followed by Trebouxiophyceae. Rotifera comprised 49.8% of the total zooplankton community (individuals L^?1), the largest proportion of any group. Temporal variations impacted the plankton abundance and community structure, and plankton abundance were more abundant during summer. The pond age did not influence the phytoplankton abundance, whereas zooplankton abundance was higher in older ponds.     

Seasonal fluctuation in the zooplankton community of Azibo reservoir (Portugal)

The seasonal fluctuation in the zooplankton community of Azibo reservoir (Portugal), was studied from November 1986 till November 1987.Fifty-three species of Protozoa, Rotifera, Copepoda and Cladocera were found. The zooplankton community was numerically dominated by rotifers, which represented 66% of the total.The highest rotifer density was reached in the beginning of spring. It decreased after the cladocerans peaked in May.Copepods had two maxima, one in April and another one in September.Although protozoans rose to high densities during winter (1.07 × 10⁵ ind m^–3), their contribution to total biomass was small (2.03%).     

浙江饮用水源地浮游动物群落特征及环境响应

2010年1月至2011年10月,对浙江省16个饮用水源地(H1—H8为河网型,K1—K8为水库型)的浮游动物群落进行季节调查。共记录浮游动物优势种(属)21种(轮虫8种、枝角类5种、剑水蚤5种和哲水蚤3种),各类群第一优势种分别为针簇多肢轮虫(Polyarthra trigla)、长额象鼻溞(Bosmina longirostris)、粗壮温剑水蚤(Thermocyclops dybowskii)和汤匙华哲水蚤(Sinocalanus dorrii)。2年间,河网和水库浮游动物平均密度分别为345.2 L-1和199.4 L-1,生物量分别为0.667 mg/L和0.421mg/L。各类群密度百分比例均以轮虫和桡足类无节幼体为主,甲壳动物以剑水蚤为主。经逐步回归分析表明,浮游动物群落密度(生物量)与河网水质因子(P0.01)之间相关性比水库(P0.05)更密切,总磷和氨氮分别入选了河网和水库所有有效的回归方程中。通径分析和决策系统分析表明,河网的总磷和叶绿素a含量对浮游动物群落变动具有正效应,溶解氧具有负效应;总磷含量是影响河网群落变动的最重要因子,叶绿素a含量则是影响群落增长最主要的限制因子。河网剑水蚤、无节幼体和轮虫群落的密度(生物量)与水体综合营养指数TLIc密度(TLIc生物量)之间有显著的线性回归关系(P0.001),无节幼体密度构成了TLIc密度变动的限制因子,轮虫生物量成为TLIc生物量变动的限制因子,而剑水蚤是一类最重要、稳定的水质指示群落,这对于筛选浮游动物群落的一些拓展性监测指标具有重要的参考作用。     

A Short-Term Study of Vertical and Horizontal Distribution of Zooplankton During Thermal Stratification in Lake Kinneret,Israel

This study documents for the first time both vertical and horizontal distribution patterns of the zooplankton community in Lake Kinneret during the period of thermal stratification. The zooplankton distribution patterns were explored in relation to abiotic (temperature, oxygen) and biotic (picocyanobacteria, ciliates, flagellates, phytoplankton, fish) environmental gradients. Sampling was carried out on 6–7 July 1992 at five stations and six depths from nearshore to offshore. Zooplankton abundance and biomass varied from 5 to 267 ind. l^–1(mean: 95 ind. l^–1), and from 0.1 to 65 d.w. mg m^–3(mean: 24 d.w. mg m^–3). Zooplankton taxonomic groups (Rotifera, Cladocera, Cyclopoida, Calanoida) and size classes (micro-, meso- and macrozooplankton) showed peaks of maximal density and biomass in the epilimnetic and metalimnetic strata (5 and 14 m). Depth, accounting for 31–39% of total spatial variation, reflected the vertical distribution of zooplankton in relation to temperature and oxygen declines, and the higher concentration of food resources (protists and phytoplankton) in the epilimnion and metalimnion. Onshore–offshore distance, accounting for 17–22% of the total spatial variance, reflected different distribution patterns shown among zooplankton groups and size classes. The macrozooplankton (Copepoda, Cladocera) was more abundant offshore, whereas microzooplankton (Rotifera and nauplii) predominated nearshore. These horizontal distribution patterns were related to small increases in temperature and phytoplankton biomass, and higher concentrations of fish in the littoral zone. Although limited to a short temporal scale, our study indicated that zooplankton spatial distribution in Lake Kinneret during the period of thermal stratification was related to physicochemical, food and predation factors, manifested differently along the vertical and nearshore–offshore gradients.     

Nitrogen removal from secondary effluent by using integrated constructed wetland system

The treatment capacity of an integrated constructed wetland system (CWS) that was designed to reduce nitrogen (N) from secondary effluent was explored. The integrated CWS consisted of vertical-flow constructed wetland, floating bed and sand filter. The vertical-flow wetland was filled with gravel, steel slag and peat from the bottom to the top. Vetiver zizanioides was selected to grow in the vertical-flow constructed wetland and Coix lacrymajobi L. was grown in the floating bed. The results showed that the integrated CWS displayed superior removal efficiency for nitrate nitrogen (NO₃⁻-N), ammonia nitrogen (NH₄⁺-N), nitrite nitrogen (NO₂⁻-N), and total nitrogen (TN). The average NO₃⁻-N, NO₂⁻-N, NH₄⁺-N and TN removal efficiencies of the integrated CWS were 98.83%, 95.60%, 98.05% and 92.41%, respectively, during the whole experimental operation. The integrated CWS may have a good potential for removing N from secondary effluent.     

Planktonic food chains of a highly humic lake

The development and metabolism of epilimnetic plankton from a highly humic lake was followed in late summer, when the predominant zooplankton species, Daphnia longispina, was very abundant (ca. 200 ind. l^?1). The experiment was made in two tanks: one with an unaltered plankton assemblage and one with larger zooplankton removed. The scarce phytoplankton community was also simple, consisting mainly of one Cryptomonas and two Mallomonas species. The abundance and species composition of smaller plankton was heavily influenced by grazing of Daphnia. In particular, the biomass, of heterotrophic flagellates increased after the removal of Daphnia. The biomass and production of bacterioplankton were not affected, and remained several times higher than that of phytoplankton. Bacterial production and grazing on bacteria were balanced, and when Daphnia was removed its grazing activity was compensated by flagellates. The removal of Daphnia did not affect the respiration or community net production of plankton. Among organisms smaller than zooplankton, bacteria seemed to be responsible for most of the respiration. The community net production was consistently negative even at the water surface, indicating an allochthonous carbon source. The results suggest that phytoplankton primary production was insufficient for the secondary production in the epilimnetic water of the study lake. The food requirements of bacteria and zooplankton, as well as of flagellates, each exceeded that supplied by phytoplankton primary production. The simple food chains in this experiment made it possible to reveal the functioning of the community so completely that dissolved organic matter is certainly comparable to or exceeds the importance of phytoplankton primary production as an energy and carbon source for food webs in this humic lake.     

Relationship between epipelon,epiphyton and phytoplankton in two limnological phases in a shallow tropical reservoir with high Nymphaea coverage

Macrophytes and phytoplankton are recognized as having roles in determining alternative stable states in shallow lakes and reservoirs, while the role of periphyton has been poorly investigated. Temporal and spatial variation of phytoplankton, epipelon and epiphyton was examined in a shallow reservoir with high abundance of aquatic macrophytes. The relationships between algae communities and abiotic factors, macrophyte coverage and zooplankton density were also analyzed. Monthly sampling was performed in three zones of the depth gradient of the reservoir. Two phases of algal dominance were found: a phytoplankton phase and epipelon phase. The phase of phytoplankton dominance was characterized by high macrophyte coverage. Rotifera was the dominant zooplankton group in all the zones. Flagellate algae were dominant in phytoplankton, epipelon and epiphyton. Macrophyte coverage was found to be a predictor for algal biomass. Changes in biomass and species composition were associated with macrophyte cover variation, mainly the Nymphaea. In addition to the abiotic factors, the macrophyte coverage was a determining factor for changes to the algal community, contributing to the alternation between dominance phases of phytoplankton and epipelon. The macrophyte–phytoplankton–periphyton relationship needs to be further known in shallow reservoirs, especially the role of epipelon as an alternate stable state.

     

Water quality and plankton periodicity in two contrasting mine lakes in Jos,Nigeria

Water quality and plankton periodicity was studied in two mine lakes near Jos in the younger granite area of Plateau State, Nigeria. The investigation was carried out for 8 months. Transparency, pH and NO₃-N were significantly higher in Lake II while DOM, alkalinity, PO ₄ ^3– -P, BOD and chloride were significantly higher in Lake I. The order of dominance in Lake I was Bacillariophyceae, Cyanophyceae, Chlorophyceae and Dinophyceae while for Lake II; Chlorophyceae, Cyanophyceae, Bacillariophyceae and Dinophyceae. In Lake I the zooplankton in order of dominance were Cladocera, Rotifera, Copepoda, and nauplii and in Lake II Cladocera, Copepoda, Rotifera and nauplii. Although both lakes would seem unproductive based on the PO ₄ ^3– -P and NO ₃ ^– -N levels, Lake I appears more productive than Lake II.     

2010年夏季雷州半岛海岸带浮游植物群落结构特征及其与主要环境因子的关系

2010年夏季对雷州半岛海岸带的浮游植物群落结构及相关环境因子进行了调查,结果共检出浮游植物211种(含变种、变型),隶属于7门76属;优势种13种,以广温、广盐沿岸性种和温带沿岸性种为主,群落结构具有亚热带和沿岸性特征。半岛海岸带浮游植物平均种类数和细胞丰度的平面分布,均由近岸往远岸逐渐减少,此外种类数是西南部>西北部>南部>东南部>东北部,细胞丰度则东北部>南部>西南部>西北部>东南部。除了东北部海岸带,多样性指数和均匀度平均值分别低于3及0.7外,其他海岸带均高于此值,物种非常丰富。相关分析表明:浮游植物种类数与水温、Si/P呈极显著的正相关,与SiO23--Si、Si/N呈显著的正相关,与NO-2-N、PO34--P呈极显著的负相关,与盐度、pH值等不存在统计学意义上的相关性;浮游植物细胞丰度与pH值、叶绿素a、DO、NO-2-N呈极显著的正相关,与盐度呈极显著的负相关,与水温、NO-3-N等不存在统计学意义上的相关性;浮游植物种类数与细胞丰度之间也不存在统计学意义上的相关性。根据浮游植物生物学指标对雷州半岛海岸带的富营养化程度进行现状评价表明:雷州半岛海岸带水质营养类型属于中-贫营养型,而根据营养盐指标评价雷州半岛海岸带水质则总体上为氮限制中度营养或氮限制潜在性富营养化水平。     

Pheopigment dynamics,zooplankton grazing rates and the autumnal ammonium peak in a Mediterranean lagoon

The dilution technique was used to estimate chlorophyll and pheopigment, net and gross production as well as zooplankton grazing over a 12-month period in a coastal lagoon in Southern France. Chlorophyll a (Cha) based gross growth rates of phytoplankton ranged from undetectable in February to 2.6 day⁻¹ in June, corresponding to 3.8 divisions per day. Cha-based grazing rates ranged from undetectable in February to 1.1 d⁻¹ in June. The seasonal growth pattern of picoplankton was similar to that of the whole community, with a peak in July, corresponding to four divisions per day. Grazing processes represented from 20 to 150% of the phytoplankton daily growth, and the grazing pressure was stronger on small phytoplankton cells than on larger cells. Gross growth rates of phytoplankton were related to zooplankton grazing rates, and both were related to water temperature. Mesozooplankton which escaped sampling or oysters had to be also invoked as additional sinks for the primary production. In the fall, pheopigment concentrations greater than chlorophyll concentrations coincided with high ammonium levels in the water column. Pheopigment a production rates were highly correlated to chlorophyll -based microzooplankton grazing rates. The pheopigment a to chlorophyll a ratio was correlated with ammonium concentrations and could be used an index of the balance between ammonium supply (degradation) and demand (uptake by phytoplankton). In addition, pheopigment degradation rates in absence of grazing could be related to irradiance, indicating photo-degradation of these compounds.     

Fish-zooplankton interactions and their effects on water quality of a tropical Brazilian reservoir

The influence of zooplanktivorous fishes on the plankton community and water quality of Americana Reservoir, Brazil was studied experimentally in 4 floating enclosures during the dry seasons (July–September) of 1982 and 1983. Two enclosures were stocked with adult fish (Astyanax bimaculatus in 1982;A. fasciatus in 1983) at near maximal densities measured in the reservoir upper surface waters (35 m^–2) and two were fish-free during each experiment lasting about one month. Marked differences were evident between the fish and fish-free enclosures after a 2–3 week period in each experiment, particularly with respect to water transparency, phytoplankton biomass, and zooplankton abundance as well as species and size composition. By the end of each experiment water transparencies were lower and phytoplankton biomass higher in the fish enclosures compared to those without fish. Also at that time Rotifera were the prominent zooplankters in the fish enclosures and Cladocera in the fish-free ones. Larger or more conspicuous species of Cladocera asDaphnia gessneri, D. ambigua, andMoina micrura were present in the fish-free enclosures but not in the fish enclosures. The interactions between fish predation, zooplankton grazing, phytoplankton biomass and water quality conditions are discussed in relation to eutrophication of a tropical aquatic ecosystem.     

Plankton communities of an acidic,polymictic, brown-water lake

The plankton of an acidic, polymictic, brown-water lake was investigated over a one-year period. The phytoplankton community was dominated by Chlamydomonas sp., Melosira varians and Peridinium pusillum during different times of the year. Densities were abnormally low, with a maximum of only 13,781 individuals per liter during the March bloom. The water column was nearly void of phytoplankton during the fall. These low densities were attributed to several of the physical and chemical characteristics of the water, including low pH, low levels of nutrients and light limitation. The latter factor was important because of low light transmission into the highly colored water, shading from macrophytes and the nearly continuous transport of the phytoplankton into the aphotic zone due to mixing of the water column by sea breezes.The zooplankton was dominated by Diaptomus floridanus, Polyarthra vulgaris, Keratella cochlearis and Daphnia ambigua. The observed densities were typical of southeastern oligotrophic lakes. The zooplankton community did not exhibit a decrease in density that corresponded with the observed low phytoplankton densities, suggesting the possibility that they relied heavily on bacterial and detrital food sources.     

Effectiveness of phytoplankton control by large-bodied and small-bodied zooplankton

Employingin situ enclosures containing inocula of the lake zooplankton (mainlyDaphnia galeata, Daphnia cucullata andBosmina spp.) from a moderately eutrophic Lake Ros (Northern Poland) or large-bodiedDaphina magna, the following observations on succession of phytoplankton were made: 1) whereasD. magna could control the density of all the photoplankton size classes, the lake zooplankton could not suppress the large-sized phytoplankters or net phytoplankton; 2) the lake zooplankton was able to control the density of small algae (< 50μm), but its effect on large algae may be opposite: a promotion of net phytoplankton growth by removing small-sized algae which can out-compete net phytoplankton for limited PO₄-P resources (<5μg P l⁻¹). Since efficiency of phytoplankton density control byD. magna decreased with an increase in net phytoplankton abundance, biomanipulation could not be successful without introducing or maintaining a high population of large-bodied cladoceran species before high densities of large algae would make the control of phytoplankton inefficient.     

Contemporary state of the zooplankton in Lake Peipsi

L. Peipsi is one of the richest fish lakes in Europe. Planktivorous smelt dominates in the fish fauna. The abundance of zooplankton fluctuates between 43 600–2241 500 ind m^–3, with the average 974 000 ind m^–3, biomass ranges from 0,09–3,69 g m^–3, with the average 1,86 g m^–3. Since the 1960s the abundance of rotifers has risen considerably while the mean zooplankter weight (B/N) has decreased from 0.005 mg to 0.004 mg. Zooplankton production (herbivores 20.6, predators 1.8, whole zooplankton community 22.4 g C m^–2 per period between May and October) can be considered high. Predatory zooplankton eats on an average 50% of the production of herbivorous zooplankton; about 50% of the whole zooplankton production (P_{Filt + Pred}) reaches fishes. The production of herbivorous zooplankton constitutes 10.1% of primary production. This ratio indicates a direct relationship between zoo- and phytoplankton in the food chain; the detrital food chain seems of little importance. About 6% of phytoplankton energy reaches fishes. The transformation of energy in the food web is efficient. On the basis of zooplankton L. Peipsi can be considered a moderately eutrophic or meso-eutrophic lake.     

Impact of submerged macrophytes on fish-zooplanl phytoplankton interactions: large-scale enclosure experiments in a shallow eutrophic lake

1. The impact of changes in submerged macrophyte abundance on fish-zooplankton-phytoplankton interactions was studied in eighteen large-scale (100 m²) enclosures in a shallow eutrophic take. The submerged macrophytes comprised Potamategon pectinatus L., P. pusillus L. and Callitriche hermaphroditica L. while the fish fry stock comprised three-spined sticklebacks, Gasterosteus acuteatus L., and roach, Rutilus rutilus L. 2. In the absence of macrophytes zooplankton biomass was low and dominated by cyclopoid copepods regardless of fish density, while the phytoplankton biovolume was high (up to 38 mm³1) and dominated by small pennate diatoms and chlorococcales. When the lake volume infested by submerged macrophytes (PVI) exceeded 15–20% and the fish density was below a catch per unit effort (CPUE) of 10 (approx. 2 fry m^?2), planktonic cladoceran biomass was high and dominated by relatively large-sized specimens, while the phytoplankton biovolume was low and dominated by small fast-growing flagellates. At higher fish densities, zooplankton biomass and average biomass of cladocerans decreased and a shift to cyclopoids occurred, while phytoplankton biovolume increased markedly and became dominated by cyanophytes and dinoflagellates. 3. Stepwise multiple linear regressions on log-transformed data revealed that the biomass of Daphnia, Bosmina, Ceriodaphmia and Chydorus were all significantly positively related to PVI and negatively to the abundance of fish or PVI x fish. The average individual biomass of cladocerans was negatively related to fish, but unrelated to PVI. Calculated zooplankton grazing pressure on phytoplankton was positively related to PVI and negatively to PVI x fish. Accordingly the phytoplankton biovolume was negatively related to PVI and to PVI x zooplankton biomass. Cyanophytes and chryptophytes (% of biomass) were positively and Chlorococcales and diatoms negatively related to PVI, while cyanophytes and Chlorococcales were negatively related to PVI x zooplankton biomass. In contrast diatoms and cryptophytes were positively related to the zooplankton biomass or PVI x zooplankton. 4. The results suggest that fish predation has less impact on the zooplankton community in the more structured environment of macrophyte beds, particularly when the PVI exceeds 15–20%. They further suggest that the refuge capacity of macrophytes decreases markedly with increasing fish density (in our study above approximately 10 CPUE). Provided that the density of planktivorous fish is not high, even small improvements in submerged macrophyte abundance may have a substantial positive impact on the zooplankton, leading to a lower phytoplankton biovolume and higher water transparency. However, at high fish densities the refuge effect seems low and no major zooplankton mediated effects of enhanced growth of macrophytes are to be expected.