Nutrients,light and primary production by phytoplankton and microphytobenthos in the eutrophic,turbid Westerschelde estuary (The Netherlands)

Abiotic factors and primary production by phytoplankton and microphytobenthos was studied in the turbid Westeschelde estuary. Because of the high turbidity and high nutrient concentrations primary production by phytoplankton is light-limited. In the inner and central parts of the estuary maximum rates of primary production were therefore measured during the summer, whereas in the more marine part spring and autumn bloom were observed. Organic loading is high, causing near anaerobic conditions upstream in the river Schelde. Because of this there were no important phytoplankton grazers in this part of the estuary and hence the grazing pressure on phytoplankton was minimal. As this reduced losses, biomass is maximal in the river Schelde, despite the very low growth rates.On a number of occasions, primary production by benthic micro-algae on intertidal flats was studied. Comparison of their rates of primary production to phytoplankton production in the same period led to the conclusion that the contribution to total primary production by benthic algae was small. The main reason for this is that the photosynthetic activity declines rapidly after the flats emerged from the water. It is argued that CO₂-limitation could only be partially responsible for the noticed decrease in activity.     

Summer feeding activities of zooplankton in Prydz Bay, Antarctica

Grazing of dominant zooplankton copepods (Calanoides acutus, and Metridia gerlachei), salps (Salpa thompsoni) and microzooplankton was determined during the austral summer of 1998/1999 at the seasonal ice zone of the Prydz Bay region. The objective was to measure the ingestion rates of zooplankton at the seasonal ice zone, so as to evaluate the importance of different groups of zooplankton in their grazing impact on phytoplankton standing stock and primary production. Grazing by copepods was low, and accounted for <1% of phytoplankton standing stocks and 3.8-12.5% of primary production for both species during this study; even the ingestion rates of individuals were at a high level compared with previous reports. S. thompsoni exhibited a relatively high grazing impact on primary production (72%) in the north of our investigation area. The highest grazing impact on phytoplankton was exerted by microzooplankton during this investigation, and accounted for 10-65% of the standing stock of phytoplankton and 34-100% of potential daily primary production. We concluded that microzooplankton was the dominant phytoplankton consumer in this study area. Salps also played an important role in control of phytoplankton where swarming occurred. The grazing of copepods had a relatively small effect on phytoplankton biomass development.     

太湖湖岸带浮游植物初级生产力特征及影响因素

利用高频溶解氧监测,估算了太湖梅梁湾湖岸带浮游植物初级生产力的高频变化特征。结合同步气象监测及浮游植物、浮游动物和营养盐的周年逐周观测数据,分析了气象和环境条件对富营养化湖泊浮游植物初级生产力的影响。结果发现,高频溶解氧监测估算的初级生产力变化与浮游藻类生物量的变化一致,能够反映出浮游植物生产力的昼夜变化、季节变化等规律。统计分析表明,气温对太湖这一富营养化湖泊初级生产力影响很大;氮的供给与浮游动物的选择性牧食也是影响浮游植物初级生产力的重要因素。湖岸带的水华堆积过程对初级生产力影响巨大,气象、水文过程又加剧了蓝藻水华初级生产力的变化幅度,反映出富营养化湖泊初级生产力可能存在极大的时空不均一性。研究表明,溶解氧高频监测法估算初级生产力能够捕捉到湖泊初级生产力的快速变化过程,可以用于富营养化湖泊初级生产力监测、蓝藻水华灾害预警中。     

Effects of sediment resuspension on phytoplankton production: teasing apart the influences of light, nutrients and algal entrainment

1. Wind‐induced sediment resuspension can affect planktonic primary productivity by influencing light penetration and nutrient availability, and by contributing meroplankton (algae resuspended from the lake bed) to the water column. We established relationships between sediment resuspension, light and nutrient availability to phytoplankton in a shallow lake on four occasions. 2. The effects of additions of surficial sediments and nutrients on the productivity of phytoplankton communities were measured in 300 mL gas‐tight bottles attached to rotating plankton wheels and exposed to a light gradient, in 24 h incubations at in situ temperatures. 3. While sediment resuspension always increased primary productivity, resuspension released phytoplankton from nutrient limitation in only two of the four experiments because the amount of available nitrogen and phosphorus entrained from the sediments was small compared with typical baseline levels in the water column. In contrast, chlorophyll a entrainment was substantial compared with baseline water column concentrations and the contribution of meroplankton to primary production was important at times, especially when seasonal irradiance in the lake was high. 4. Comparison of the in situ light climate with the threshold of light‐limitation of the phytoplankton indicated that phytoplankton in the lake were only likely to be light‐limited at times of extreme turbidity (e.g. >200 nephelometric turbidity units), particularly when these occur in winter. Therefore, resuspension influenced phytoplankton production mainly via effects on available nutrients and by entraining algae. The importance of each of these varied in time. 5. The partitioning of primary productivity between the water column and sediments in shallow lakes greatly influences the outcome of resuspension events for water column primary productivity.     

Spatio-temporal study of phytoplankton cell viability in a eutrophic reservoir using SYTOX Green nucleic acid stain

Despite the global importance of phytoplankton primary production, the ecological role of cell death as an important loss process in phytoplankton is poorly understood. To assess the significance of cell death in phytoplankton, we studied cell viability of dominant species in the canyon-shaped eutrophic ?ímov Reservoir (Czech Republic) at weekly and biweekly intervals from April to October 2011. Surface samples were taken from the lacustrine zone (near the dam, low nutrient level) and transition zone (near the river inflow, high nutrient level) of the reservoir. Moreover, samples from euphotic depth (1% of surface irradiance) were taken from the lacustrine zone. We used the membrane-impermeant nucleic acid dye SYTOX Green to examine seasonal and spatial differences in phytoplankton cell viability. Three species (diatoms Asterionella formosa, Fragilaria crotonensis, and cyanobacterium Aphanizomenon flos-aquae) were studied in detail. There was no difference in Asterionella cell viability among sampling sites. In the lacustrine zone, Fragilaria and Aphanizomenon exhibited lower viability than in the transition zone. In addition, Aphanizomenon viability was significantly lower at the euphotic depth. Nutrient levels were revealed as a factor influencing Fragilaria viability, while light availability was more important for Aphanizomenon. Our results evidenced that the importance of cell death, in particular phytoplankton taxa, varies both spatially and temporally. Moreover, our study indicates that coexisting taxa may differ in their capacity to cope with different environmental stressors.     

春季赤潮频发期东海微型浮游动物摄食研究

2002年4～5月在东海长江口及其邻近水域的8、11、14、23和28号5个典型站位采样。用现场稀释法对春季东海水域浮游植物的生长率和微型浮游动物对浮游植物的摄食压力等方面进行了研究．结果表明,微型浮游动物的摄食行为在东海赤潮过程起到关键作用．各站位微型浮游动物主要以急游虫、红色中缢虫和夜光藻为主,在种类上砂壳纤毛虫是主要的类群．微型浮游动物的摄食速率范围在0．28～1．13d-1,对浮游植物现存量的摄食压力范围在35．14％～811．69％。对浮游植物潜在初级生产力的摄食压力范围在74．04％～203．25％,对浮游植物碳的摄食率范围在9．58～97．91μg·L-1·d-1,靠近岸边的站位,微型浮游动物的摄食速率、对浮游植物现存量的摄食压力和对浮游植物碳的摄食率相对较高。而远离岸边的站位对浮游植物潜在初级生产力的摄食压力却较高．与世界其它海区比较此水域微型浮游动物摄食压力处于较高水平．急游虫是控制东海主要赤潮原因生物具齿原甲藻生长的关键种类．     

The physical-chemical limnology of a desert reservoir

The physical-chemical limnology of a desert lake in central Arizona was investigated from February 1971–July 1973. The reservoir was found to be a warm-monomictic, eutrophic lake which exhibited pronounced thermal stratification during the summer months. Surface water temperatures ranged from a minimum of ca. 9 C in January to a maximum near 30 C in July–August. Chemically the lake may be described as a hard water lake of moderately high alkalinity and salinity. The concentration of the principal ions was affected significantly by precipitation run-off. The primary nutrients N and P were subject to considerable seasonal variability, being influenced most by precipitation and phytoplankton abundance. Trace element concentrations were low and showed comparatively little fluctuation during the study.     

Phytoplankton growth rates in the freshwater tidal reaches of the Schelde estuary (Belgium) estimated using a simple light-limited primary production model

During the course of 1996, phytoplankton was monitored in the turbid, freshwater tidal reaches of the Schelde estuary. Using a simple light-limited primary production model, phytoplankton growth rates were estimated to evaluate whether phytoplankton could attain net positive growth rates and whether growth rates were high enough for a bloom to develop. Two phytoplankton blooms were observed in the freshwater tidal reaches. The first bloom occurred in March and was mainly situated in the most upstream reaches of the freshwater tidal zone, suggesting that it was imported from the tributary river Schelde. The second bloom occurred in July and August. This summer bloom was situated more downstream in the freshwater tidal reaches and appeared to have developed within the estuary. A comparison between phytoplankton growth rates estimated using a simple primary production model and flushing rate of the water indicated that no net increase in phytoplankton biomass was possible in March while phytoplankton could theoretically increase its biomass by 20% per day during summer. Chlorophyllaconcentrations at all times decreased strongly at salinities between 5–10 psu. This decline was ascribed to a combination of salinity stress and light limitation. Phytoplankton biomass and estimated annual net production were much higher in the freshwater tidal zone compared to the brackish reaches of the estuary (salinity > 10 psu) despite mixing depth to euphotic depth ratios being similar. Possible reasons for this high production include high nutrient concentrations, low zooplankton grazing pressure and import of phytoplankton blooms from the tributary rivers.     

An experimental investigation of phytoplankton nutrient limitation in two contrasting low arctic lakes

We investigated whether phytoplankton communities in two lakes in SW Greenland were phosphorus or nitrogen limited. The study lakes have contrasting water chemistry (mean conductivities differ ten fold) and are located near Kangerlussuaq, SW Greenland (~67°N, 51°W). A microcosm nutrient enrichment experiment was performed in June 2003 to determine whether nitrate or phosphate addition stimulated phytoplankton growth. Samples were analysed for species composition, biomass, and alkaline phosphatase activity (APA). Initially, both lakes had extremely low total phosphorus but high total nitrogen concentrations and high APA, suggesting that the phytoplankton were phosphorus limited prior to the start of the experiment. The phytoplankton composition and biomass (mainly Ochromonas spp.) responded to phosphate but not to nitrate addition. In both lakes, chlorophyll a increased significantly when phosphate was added. Furthermore, APA was significantly lower in the two lakes when phosphate was added compared to the control and the nitrogen addition treatment. The dominance of mixotrophic phytoplankton and high DOC values suggest that these lakes may be regulated by microbial loop processes.     

Contrasting responses of intertidal microphytobenthos and phytoplankton biomass and taxonomic composition to the nutrient loads in the Jiulong River Estuary

Microphytobenthos (MPB) and phytoplankton are important primary producers in the estuarial ecosystem, and their functions are critical to the ecosystem's biodiversity and environmental safety. The aim of this study was to compare the response of MPB and phytoplankton to the nutrient loads in a eutrophic estuary, which has seldom been studied. We used high‐performance liquid chromatography (HPLC) and CHEMTAX software to examine the biomass and taxonomic composition of both MPB and phytoplankton at Da‐yu Island (DYI) and Ji‐yu Island (JYI) in the Jiulong River Estuary from July 2010 to March 2012. The results showed that MPB chlorophyll a was low in the summer and high in the winter at both DYI and JYI, indicating a unimodal pattern. However, the phytoplankton chlorophyll a showed a mirrored pattern. Diatoms were the dominant class in both benthic and pelagic environments. Although redundancy analysis indicated that the effects of different environmental factors could not be easily separated, it is likely that phosphate and temperature were the most important factors regulating the seasonal patterns of MPB and phytoplankton diatoms, respectively. MPB and phytoplankton cyanobacteria was co‐limited by salinity and temperature. The high N/P ratio and low phosphate favored chlorophytes and cyanobacteria. Our study demonstrates the use of HPLC and CHEMTAX in an integrated survey of the spatial and temporal distribution patterns of MPB and phytoplankton in an estuarial ecosystem. The contrasting responses of MPB and phytoplankton to nutrient loads indicate the critical role of MPB in subtropical estuarial ecosystem function. The relationship between nutrients and MPB may indicate a significant contribution to carbon and nutrient cycling.     

Nutrients and phytoplankton primary production in the marine tidal Oosterschelde estuary (The Netherlands)

In a shallow marine tidal area, the eastern part of Oosterschelde estuary in the S.W. Netherlands, phytoplankton primary production amounted to 176–338 g C.m^?2.y^?1 during the period 1981–1985. The influence of nutrient concentrations on the phytoplankton primary production is discussed. Phosphate and inorganic nitrogen generally were amply available. Import of inorganic nitrogen into the basin was shown and an intense delivery of ammonia by zoobenthos was suggested. Nitrate was considered to be slightly influenced by phytoplankton consumption and mainly by nitrate reduction at the bottom. Silicate may have played a limiting role in phytoplankton primary production. The first phytoplankton bloom in spring (diatom bloom) always terminated when silicate concentration decreased below K_s values. Further on in 1983 and 1984 both primary production and chlorophyll curves showed a dip when silicate was not available. The influence of available light on the primary production was demonstrated during situations with a low extinction coefficient when primary production reached maximum values. Further on during 1985 the spring bloom occurred already in March when winter extinction coefficients were lower than during preceding winters. Long term production studies are necessary to understand the extreme fluctuations of annual production patterns in relation to the prevailing environmental conditions.     

The plankton ecology of Lake St. Clair, 1984

Lake St. Clair phytoplankton and zooplankton abundance and composition was analyzed during the period of May to September 1984. In addition, size-fractionated primary productivity and other limnological parameters were measured. Highest phytoplankton biomass was observed during spring (May) with high values for the southern and southeastern regions of the lake. Seasonally, the mean phytoplankton biomass ranged between 0.17 and 1.18 g m^-3 with high values recorded during spring (May, June) compared to summer. In the spring the phytoplankton was dominated by Diatomeae followed by Chrysophyceae and Cryptophyceae. During the summer the diatoms showed a decreasing trend due to the relative prevalence of Chrysophyceae, Cryptophyceae, and Chlorophyta. The species composition was oligotrophic-mesotrophic with mixed occurrence of some eutrophic species. The phytoplankton size composition indicated dominance of microplankton/netplankton (> 20 µm) and ultraplankton (< 20 µm) during spring and summer respectively. On an overall basis ultraplankton contributed overwhelmingly to primary productivity, as much as 75 percent in the summer.The mean zooplankton biomass ranged from 173.0 to 1306.0 mg l^- dominated by Cladocerans (bosminids) in contrast to the other Great Lakes. Statistical evaluation of the phytoplankton — nutrient-contaminant interactions revealed positive correlations with heavy metals, suggestive of a physiological adaptation to contamination from the chemical valley. Based on low biomass, high Production/Biomass ratio, dominance of ultraplankton, characteristic species composition and plankton spectra, the lake appears to be an oligotrophic-mesotrophic perturbed ecosystem.     

Phytoplankton in the surface and chlorophyll maximum off southern California in August, 1978

A synoptic sampling of 30 stations in a 25 km x40 km grid closeto the southern California coast revealed that chlorophyll andphytoplankton biomass were higher nearshore and to the south,where relatively large dinoflagellates dominated. A subsurfacechlorophyll maximum was present at each station, near the nitraclineand 10% light level. Nearshore and to the south, these maximarepresented higher levels of phytoplankton biomass than at thesurface whereas at the other stations, chlorophyll maxima couldbe attributed to physiological changes in chlorophyll contentof small flagellated phytoplankton rather than to the accumulationof phytoplankton at depth. Univariate and multivariate statisticalprocedures were used to demonstrate that phytoplankton assemblagesseparated by tens of meters in the vertical were just as differentas those separated by tens of kilometers in the horizontal.We conclude that residence time of water near the coast is ofgreat importance to the determination of the abundance and taxonomiccharacteristics of phytoplankton, and that advection of offshorewater towards the coast was the major determinant of the patternobserved during our study. ¹Present address: University of Texas, Port Aransas Marine Laboratory,Port Aransas, TX 78373, USA     

The time course of phytoplankton biomass and related limnological factors in shallow and deep lakes: a multivariate approach

Weekly studies of phytoplankton biomass and environmental variables were made over one year in a shallow stratifying, hypertrophic El Porcal lake near Madrid (Spain). Data were collected on abiotic factors, primary production, biomass and phytoplankton losses and subjected to reduction by means of several principal component analyses. Furthermore, weekly data on the same variables were gathered from published studies on Überlinger See, an embayment of the much deeper, mesotrophic Lake Constance (Central Europe), and treated in the same way. The two first principal components of PCAs on biological variables explained more than 60% of overall variance in both lakes. They could be ascribed to phytoplankton production + biomass and photosynthetic physiology + phytoplankton losses, respectively. The ordination of the biological trajectories in the data space of the two first principal components revealed six stable states of phytoplankton biomass in the shallow lake and seven in the deep lake. The breakpoints between stable states could be due to environmental, abiotic variables in some cases but biological interactions were suspected to be the cause of the other breakpoints. The abiotic effects on phytoplankton biomass took longer to occur in the deep lake. Also, short-term dynamics (one-three weeks) were demonstrated for both phytoplankton communities.A preliminary comparison between phytoplankton biomass dynamics in stratifying, shallow and deep lakes suggests that differences may be attributed partly to differences in depth.     

Attached and Free-Floating Bacterioplankton in Howe Sound, British Columbia, a Coastal Marine Fjord-Embayment

Factors which influence the attachment of bacterioplankton to particles (including phytoplankton) were investigated by using (i) water samples removed from a coastal temperate fjord over an annual cycle and (ii) unialgal cultures of Prorocentrum minimum, Dunaliella tertiolecta, and Skeletonema costatum. Silt and salinity levels in this fjord seawater did not appear to influence bacterial attachment, but the percent attached bacteria was inversely related to both chlorophyll a concentrations and primary productivities. During periods of high primary productivities the percent attached bacteria was low, whereas during periods of low, increasing, and declining primary productivities the percent attached bacteria was high. A similar pattern of bacterial attachment was observed when the three phytoplankton were grown as batch cultures. The percent attached bacterial numbers increased upon the initiation of algal growth and after these cells stopped growing, but not while the algae were growing. We suggest that a major factor influencing the attachment of bacterioplankton is the physiological condition of their major nutrient source, the phytoplankton; mainly free-living bacteria are associated with growing phytoplankton, whereas a much greater proportion of the bacteria are attached among senescent phytoplankton populations.     

Ingestion rates of phytoplankton by copepod size fractions on a bloom associated with an off-shelf front off NW Spain

Herbivory by copepods was studied from the coast towards theocean, during a bloom in May 1994 off NW Spain.Ingestion rateswere estimated by the gut chlorophyll content method in threesize fractions. The chlorophyll content displayed significantdaily cycles. Three different water bodies were described: coastal,shelf break and oceanic; the latter two zones separated by athermo haline frontal structure. Marked differences in planktonspecies composition, vertical distribution and biological rateswere found between zones. Theiiighest phytoplankton biomass,dominated by chain- forming diatoms, occurred in the oceaniczone associated with low primary production rates. Copepod feedinghad a low effect on oceanic phytoplankton; up to 0.2% of carbonstock and<3% of carbon production was consumed daily. Incontrast, medium-sized and large copepods removed 3% of carbonstock and 12% of primary production daily near the coast, wherephytoplankton were dominated by small flagellates in activegrowth. The highest variability in both plankton compositionand ingestionrates was found in the shelf-break zone, probablydue to displacements of the front. Copepods exerted a moderatepredation pressure on phytoplankton in coastal waters. Meanwhile,the impact of copepods on the offshore bloom was negligibleand the fate of the accumulated particulate carbon would bemostly determined by sedimentationand water dynamics.     

Phytoplankton distribution and production along a wide environmental gradient in the South-West Atlantic off Uruguay

We investigated phytoplankton biomass, assemblage structure and production along an environmental gradient to evaluate if chlorophyll-a (as proxy for biomass) and primary production peaked under conditions hypothesised to favour phytoplankton growth. During Spring 2003, a wide area from shallow estuarine waters to the shelf slope off the Río de la Plata was sampled and routine measurements included CTD profiles, nutrients, chlorophyll-a, phytoplankton composition and abundance, seston and organic matter loads, downwelling light and, at selected stations, production versus irradiance experiments. Spatial differences in abiotic variables suggested distinct hydrographic zones that differed in phytoplankton biomass and productivity. Chlorophyll-a was highest under estuarine influence and peaked at low salinity when strong stratification developed in the outer estuary, and was minimum at the shelf break and slope. In that area, however, relatively high chlorophyll-a was associated to oceanographic fronts and to the occurrence of Sub Antarctic water within the photic depth range. Productivity was maximum in shallow waters, but biomass-specific productivity peaked at the outer shelf in oceanographic fronts or in upwelled Sub Antarctic waters. Over shelf and slope waters productivity and biomass were not tightly coupled, as indicated by situations of high biomass and low productivity (Station 9), low biomass and high productivity (Station 10), or both high biomass and productivity (Station 22). Ordination analysis of phytoplankton taxa suggested that assemblages changed gradually along the environmental gradient and correlated to abiotic variables defining geographic zones. Overall results were consistent with an interpretation that phytoplankton biomass and growth were modulated by light in estuarine and coastal waters, and by hydrographic processes on the continental shelf and slope. Handling editor: Luigi Naselli-Flores     

Predicting the spatial dominance of phytoplankton in a light limited and incompletely mixed eutrophic water column using the PROTECH model

1. A phytoplankton community model [Phytoplankton RespOnses To Environmental CHange (PROTECH)] was used to examine the effect of a wide range of varied light intensities and mixed depths upon simulated phytoplankton populations. Two different column lengths of the simulated water body were examined (the upper 5 m and the whole 14.5 m water column) for each scenario.
2. The hypotheses tested were that: (i) under low light intensity and/or deep mixing the simulated community will be dominated by a phytoplankter with a low critical light intensity; (ii) at high light intensity and shallow mixing the simulated community will be dominated by small, fast-growing phytoplankters; (iii) under all conditions, except deep mixing, the largest proportion of phytoplankton biomass will be found near the surface.
3. It was found under most conditions that, although there was a bloom in the upper column (dominated by algae such as Chlorella , Ceratium or Rhodomonas ), the largest phytoplankton biomass in the water column was located 9 m below the surface and consisted of solely Asterionella . This bloom was missed by the 5-m samples. Thus, using the whole column sample lengths, hypothesis (i) was not rejected but hypotheses (ii) and (iii) were refuted.
4. The inclusion of specific movement characteristics of phytoplankton in the model allowed the possibility of the dominance of multiple spaces within the water column and should be included in any model-based investigation of this topic. Further, the results from the model suggest that a reduced depth of mixing creates greater environmental heterogeneity, allowing more species to persist.     

Seasonal and interannual variability in the taxonomic composition and production dynamics of phytoplankton assemblages in Crater Lake,Oregon

Taxonomic composition and production dynamics of phytoplankton assemblages in Crater Lake, Oregon, were examined during time periods between 1984 and 2000. The objectives of the study were (1) to investigate spatial and temporal patterns in species composition, chlorophyll concentration, and primary productivity relative to seasonal patterns of water circulation; (2) to explore relationships between water column chemistry and the taxonomic composition of the phytoplankton; and (3) to determine effects of primary and secondary consumers on the phytoplankton assemblage. An analysis of 690 samples obtained on 50 sampling dates from 14 depths in the water column found a total of 163 phytoplankton taxa, 134 of which were identified to genus and 101 were identified to the species or variety level of classification. Dominant species by density or biovolume included Nitzschia gracilis, Stephanodiscus hantzschii, Ankistrodesmus spiralis, Mougeotia parvula, Dinobryon sertularia, Tribonema affine, Aphanocapsa delicatissima, Synechocystis sp., Gymnodinium inversum, and Peridinium inconspicuum. When the lake was thermally stratified in late summer, some of these species exhibited a stratified vertical distribution in the water column. A cluster analysis of these data also revealed a vertical stratification of the flora from the middle of the summer through the early fall. Multivariate test statistics indicated that there was a significant relationship between the species composition of the phytoplankton and a corresponding set of chemical variables measured for samples from the water column. In this case, concentrations of total phosphorus, ammonia, total Kjeldahl nitrogen, and alkalinity were associated with interannual changes in the flora; whereas pH and concentrations of dissolved oxygen, orthophosphate, nitrate, and silicon were more closely related to spatial variation and thermal stratification. The maximum chlorophyll concentration when the lake was thermally stratified in August and September was usually between depths of 100 m and 120 m. In comparison, the depth of maximum primary production ranged from 60 m to 80 m at this time of year. Regression analysis detected a weak negative relationship between chlorophyll concentration and Secchi disk depth, a measure of lake transparency. However, interannual changes in chlorophyll concentration and the species composition of the phytoplankton could not be explained by the removal of the septic field near Rim Village or by patterns of upwelling from the deep lake. An alternative trophic hypothesis proposes that the productivity of Crater Lake is controlled primarily by long-term patterns of climatic change that regulate the supply of allochthonous nutrients.