A flow cytometric approach to assessing the environmental and physiological status of phytoplankton

Individual particle analysis using a flow cytometer (FCM) was made on natural phytoplankton communities in oligotrophic waters. Our objective was to develop an assay to yield information on the nutrient history of individual cells using FCM. Results from nutrient assays showed that both biovolume and total red fluorescence are affected by the nutrient conditions in the incubator. The light effect was measured by changes in the chlorophyll content of the cells, and after the 12 h incubation the cells seemed well adapted to the light conditions. The estimated kinetic constant for the chlorophyll synthesis averaged 1.5 x 1.0(-2) h-1, whereas the growth rate calculated from the changes in the cell numbers changed from 0.14 to greater than 2.5 doubling per day. The smallest size fraction presented the highest growth rate (greater than 2.5 doublings per day). The relationship between the total red fluorescence as estimated with the FCM and the biovolume revealed that the cells from the 2 m samples at the beginning of the experiment were probably nutrient limited. Important changes in the size of the cells under nutrient limitation were also observed. The FCM data suggest that the FCM is a valuable tool for estimating the relative growth response and nutritional state of natural phytoplankton populations.     

COMBINATION OF FLOW CYTOMETRY AND SINGLE CELL ABSORPTION SPECTROSCOPY TO STUDY THE PHYTOPLANKTON STRUCTURE AND TO CALCULATE THE CHL A SPECIFIC ABSORPTION COEFFICIENTS AT THE TAXON LEVEL1

The phytoplankton community structure of a hypertrophic lake was quantitatively determined with the aid of flow cytometry. The flow cytometry signals were calibrated to obtain cell‐specific information, such as the chl a content and the biovolume per cell. The reliability of this method was tested with laboratory cultures. The results of the phytoplankton structure in a hypertrophic lake with respect to chl distribution in the different algal groups obtained by flow cytometry were compared with the results from HPLC pigment fingerprinting. Both methods yield the percentage contribution of the different algal groups to total chl a. The chl a specific absorption coefficient of the phytoplankton (a^*_Phy) was determined via visible (VIS) spectroscopy of samples taken from a hypertrophic lake (Auensee) in 2003. The results indicated that a^*_Phy of the total cell suspension is dependent on the phytoplankton structure as well as on environmental factors. The linear relationship between a^*_Phy at 675 nm and the product of the chl a content per cell and the biovolume offered the possibility to normalize phytoplankton absorption spectra to acquire the taxon‐specific a^*_Phy. The estimated a^*_Phy (675 nm) values were used to normalize single cell absorption spectra at this wavelength to obtain the a^*_Phy between 400 and 750 nm for representatives of the major algal groups. Our measurements show that the absorption coefficient for the whole phytoplankton community varies within the season. Finally, we used the a^*_Phy and the chl a distribution to calculate the light absorption of each algal group in the hypertrophic lake.     

Structure of the Phytoplankton Community and Its Relationship to Water Quality in Donghu Lake, Wuhan, China

The phytoplankton community structure, in terms of species composition, total standing crop,and abundance of the dominant algal species, at four stations in Donghu Lake, Wuhan, China, was investigated monthly from January 1994 to December 1996. A total of 260 taxa was observed, of which Chlorophyta (106 taxa) contributed the highest portion of the total number of taxa, followed by Bacillariophyta (82 taxa) and Cyanophyta (32 taxa). The total standing crop measured by means of chlorophyll a content, cell density,and cell biovolume, as well as the abundance of the dominant species, declined in the order of Station I to Station IV. Seasonal changes of the standing crop varied greatly among the four stations. Although the cell density at the four stations showed a single peak within a year, the peak density varied from July to November, dependent on the sampling year and the station. For chlorophyll a content and cell biovolume,multiple peaks were observed at Stations I and II, but a single peak was found at Stations III and IV. The phytoplankton community structure indicated that the trophic status was the highest at Station I (most eutrophic), followed by Station II; Stations III and IV were the least trophic areas. The long-term changes in phytoplankton community structure further suggested that changes in phytoplankton community structure were correlated with water quality, and eutrophication of Donghu Lake had been aggravated since the 1950s.     

The relationship between phytoplankton biovolume and chlorophyll in a deep oligotrophic lake: decoupling in their spatial and temporal maxima

The seasonal distributions of phytoplankton biovolume and chlorophylla content were monitored for 14 months in a deep oligotrophic,high mountain lake (Redó, Pyrenees). An allometric relationshipof chlorophyll with biovolume was found throughout the periodstudied, with a correlation coefficient of 0.66. However, therelationship changed with season and the taxonomic compositionof the phytoplankton. Both parameters showed a similar seasonalpattern, but differences in space and time were observed. Thechlorophyll maximum was recorded deeper and later than thatof phytoplankton biovolume. While the biovolume maximum wasrelated to an improvement in conditions for growth (nutrientinput during column mixing periods), and reflected an increasein biomass, the chlorophyll maximum was related to changes incell pigment content, and to spatial or successional trendsin species dominance. Flagellated chrysophytes predominatedat the chlorophyll maxima. Chlorophyll content per unit of phytoplanktonbiovolume fluctuated greatly throughout the year, dependingon light intensity, temperature and phytoplankton composition.Of the main groups of phytoplankton in the lake, the dinoflagellates,which dominated the summer epilimnion phytoplankton community,recorded the lowest pigment content per biovolume (which isconsistent with their size). Higher chlorophyll contents perbiovolume were found in the deep hypolimnion and during thewinter cover period associated with small cells such as somespecies of chlorococcales chlorophytes. When flagellated chrysophyteswere predominant, a broad range of chlorophyll values per biovolumewas found and there was no significant correlation between thetwo biomass indices. These findings reaffirm the need to treatphytoplankton biomass estimates with caution, in particularwhen conducting primary production studies. While our resultsshow that changes in chlorophyll content per cell occur as aphotoacclimation response along a vertical profile, they alsopoint out a component of the successional trends which appearin a phytoplankton growth phase in a lake.     

Comparing microscopic counts and pigment analyses in 46 phytoplankton communities from lakes of different trophic state

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Structure of the Phytoplankton Community and Its Relationship to Water Quality in Donghu Lake, Wuhan, China

The phytoplankton community structure, in terms of species composition, total standing crop,and abundance of the dominant algal species, at four stations in Donghu Lake, Wuhan, China, was investigated monthly from January 1994 to December 1996. A total of 260 taxa was observed, of which Chlorophyta (106 taxa) contributed the highest portion of the total number of taxa, followed by Bacillariophyta (82 taxa)and Cyanophyta (32 taxa). The total standing crop measured by means of chlorophyll a content, cell density,and cell biovolume, as well as the abundance of the dominant species, declined in the order of Station I to Station Ⅳ. Seasonal changes of the standing crop varied greatly among the four stations. Although the cell density at the four stations showed a single peak within a year, the peak density varied from July to November, dependent on the sampling year and the station. For chlorophyll a content and cell biovolume,multiple peaks were observed at Stations Ⅰ and Ⅱ, but a single peak was found at Stations Ⅲ and Ⅳ. The phytoplankton community structure indicated that the trophic status was the highest at Station Ⅰ (most eutrophic), followed by Station Ⅱ; Stations Ⅲ and Ⅳ were the least trophic areas. The long-term changes in phytoplankton community structure further suggested that changes in phytoplankton community structure were correlated with water quality, and eutrophication of Donghu Lake had been aggravated since the 1950s.     

Effects of zebra mussels on phytoplankton and ciliates: a field mesocosm experiment

Many observational studies in North American lakes have documenteddecreases in phytoplankton abundance after the invasion of thezebra mussel (Dreissena polymorpha). However, few field experimentshave examined in detail the effect of zebra mussels on phytoplanktonabundance and species composition over an extended period. Replicatedin situ mesocosms were used to evaluate the impact of naturaldensities of zebra mussels on phytoplankton and ciliate biovolume,and algal species composition over a 5-week period in a habitatthat lacked extant mussel populations. Mussel biomass used inthe experiment was determined using a regression model basedon a data analysis that predicts zebra mussel biomass from totalphosphorus concentration. Within 1 week, zebra mussels decreasedphytoplankton biovolume by 53% and ciliate biovolume by 71%.The effect of zebra mussels on ciliate biovolume was sustainedthroughout the study. However, the effect of zebra mussels onphytoplankton abundance gradually waned over the remaining 4weeks of the experiment, such that the declining effect of zebramussels could not be explained by a shift towards less edibleand/or faster growing algal species. The mussels’ decliningcondition could help to explain the effect observed over thecourse of the experiment.     

Estimating phytoplankton carbon from microscopic counts: an application for riverine systems

Algal biomass, in addition to cell numbers, is a measure of the successful conversion of inorganic to organic carbon. Consequently, carbon is the main currency used in aquatic models and in flux and budget studies. On the other hand, microscopic observation and counts remain the only means for determining species composition and biomass, which is relevant to many aspects of aquatic ecology. In this study, we focus on the way to convert biovolume to carbon biomass for algal assemblages of two rivers, using a computerized system that records dimensions of phytoplankton (Gosselain & Hamilton, 2000). We first compare different equations found in the literature for converting algal cell volume to cellular carbon content. We then evaluate the accuracy of a biomass estimate based on less time-consuming measurements, using pre-determined biovolume values instead of measuring cells in all samples. Biovolume/carbon equations are evaluated using total phytoplankton carbon biomass determined from measured chlorophyll a. Equations established for freshwater taxa seem to provide better estimates of algal biomass in the two case studies presented here, the Rideau and Meuse rivers (Canada and Belgium, respectively) than do more numerous equations defined for marine taxa. Furthermore, equations that make a distinction between diatoms and other algae appear more appropriate than those considering all algal groups as a whole. Finally, mean values of algal biovolumes, determined using sufficient measurements of cell dimensions from representative sampling series, may prove sufficient for carbon estimates of taxa in relatively homogenous size ranges. The careful choice of appropriate volumetric shapes and taxa categories remains of prime importance to get precise results.     

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.     

<Emphasis Type="Italic">In situ</Emphasis> Quantification of Phytoplankton in Reservoirs Using a Submersible Spectrofluorometer

A submersible in situ spectrofluorometer, which permits the differentiation of four algal groups (green algae, diatoms, cryptophytes and cyanobacteria), was used for phytoplankton monitoring in five reservoirs with varying levels of eutrophication and composition of their phytoplankton communities. Data obtained in situ were compared to standard laboratory methods for phytoplankton quantification; concentration of chlorophyll a and microscopy analysis. A high correlation (r = 0.95, n = 96) between chlorophyll a levels using different methods was found in all types of phytoplankton community. Taxonomic analyses and cell counts were closely related to the ratio of algal classes measured by the in situ spectrofluorometer. The submersible device used in the study measures in a continuous mode, which is advantageous in comparison with discrete sampling. This method appears to be a good tool for water quality management and can be used in the detection of natural horizontal and vertical variability in phytoplankton communities or for the early detection of cyanobacterial blooms. The device used in this study is recommended as a screening tool that enables more effective sampling that can be focused on the localities and depths where changes in phytoplankton composition occur.     

Are algal communities driven toward maximum biomass?

In this continental-scale study, we show that in major benthic and planktonic stream habitats, algal biovolume--a proxy measure of biomass--is a unimodal function of species richness (SR). The biovolume peak is observed at intermediate to high SR in the benthos but at low richness in the phytoplankton. The unimodal nature of the biomass-diversity relationship implies that a decline in algal biomass with potential harmful effects on all higher trophic levels, from invertebrates to fish, can result from either excessive species gain or species loss, both being common consequences of human-induced habitat alterations. SR frequency distributions indicate that the most frequent richness is habitat-specific and significantly higher in the benthos than in the plankton. In all studied stream environments, the most frequent SR is lower than the SR that yields the highest biovolume, probably as a result of anthropogenic influences, but always within one standard deviation from it, i.e. they are statistically indistinguishable. This suggests that algal communities may be driven toward maximum biomass.     

Phytoplankton distribution along trophic gradients within and among reservoirs in Catalonia (Spain)

1. Longitudinal gradients in the epilimnetic waters of stratified reservoirs provide a useful database to study changing environmental conditions. The spatial distribution, assemblage structure and specific adaptations of phytoplankton assemblages can be analysed along these gradients over short time scales. 2. Four reservoirs with a similar typology, located along an altitudinal gradient in the same eco‐region, were sampled along their longitudinal axes. In total, 19 sampling stations provided a trophic spectrum, ranging from oligo‐mesotrophy to hypertrophy, which was quantified by calculating the trophic state index of each sampling station in the four reservoirs. 3. Several patterns in phytoplankton assemblage structure were detected. Total chlorophyll‐a (Chl‐a), biovolume, abundance and the relative biomass contribution of the main algal groups (chlorophytes, cyanobacteria, cryptophytes and diatoms) were highly correlated with their location along the trophic gradient. 4. We also adopted the functional classification of Reynolds et al. (2002) : this effectively summarized differences among phytoplankton assemblages under varying resource‐limiting combinations, especially nutrients and underwater light climate. 5. In terms of relationships with the trophic gradient, diatoms and cyanobacteria exhibited significant opposing trends in both their relative chlorophyll contribution to total Chl‐a and biovolume. Chlorophytes were more abundant at an intermediate position along the trophic spectrum. 6. The identified patterns are consistent with models of self‐organization of phytoplankton assemblages. In particular, light availability was a strong determinant of size and shape diversity, especially in hypertrophic conditions, where ‘R‐strategist’, needle shaped species, dominated the system. In contrast, under decreased availability of nutrients and higher light extinction coefficients (K_d), the system was co‐dominated by C‐ and S‐strategist species, having shapes with a higher surface/volume ratio.     

Effects of invasive zebra mussels on phytoplankton,turbidity, and dissolved nutrients in reservoirs

Few experiments have quantified the effects of invasive zebra mussels (Dreissena polymorpha) on man-made reservoirs relative to other aquatic habitats. Reservoirs, however, are the dominate water body type in many of the states that are at the current front of the zebra mussel invasion into the western United States. The objective of this research, therefore, was to determine how zebra mussels affected phytoplankton, turbidity, and dissolved nutrients in water that was collected from three Kansas reservoirs that varied in trophic state (mesotrophic to hypereutrophic), but all experienced frequent cyanobacterial blooms. Laboratory mesocosm experiments were conducted to document the effects of zebra mussels on cyanobacteria and general water quality characteristics in the reservoir water. Zebra mussels significantly reduced algal biomass, and the total biovolume of cyanobacteria (communities were dominated by Anabaena) in each reservoir experiment. The effects of zebra mussels on other major algal groups (diatoms, flagellates, and green algae) and algal diversity were less consistent and varied between the three reservoir experiments. Similarly, the effects of zebra mussels on nutrient concentrations varied between experiments. Zebra mussels increased dissolved phosphorus concentrations in two of the reservoir experiments, but there was no effect of zebra mussels on dissolved phosphorus in the mesotrophic reservoir experiment. Combined, our results strongly suggest that zebra mussels have the potential to significantly impact reservoirs as they continue to expand throughout the western United States. Moreover, the magnitude of these effects may be context dependent and vary depending on the trophic state and/or resident phytoplankton communities of individual reservoirs as has similarly been reported for natural lakes.     

Pelagic zonation of water quality and phytoplankton in the Great Lakes

Analysis of aquatic ecosystem data collected from large water bodies must consider spatial variations. A suite of pelagic survey stations exists for the Laurentian Great Lakes, but little is known about their redundancy. We present a strategy to delineate the lakes into zones based on water quality and phytoplankton biovolume. Water samples were collected from 72 sites in two seasons (spring and summer) from 2007 to 2010 in all five lakes. Integrated samples were analyzed for phytoplankton biovolume and nine water quality parameters. We conducted cluster analysis, principal components analysis and non-metric multidimensional scaling methods for water quality and phytoplankton taxon-specific biovolume for the Great Lakes basin and for each lake separately. There were significant lake-to-lake differences, and based on lake-specific analyses, Lake Superior, Lake Michigan and Lake Erie were each divided into three zones; Lake Huron and Lake Ontario were each grouped into two zones. The zones identified by water quality and phytoplankton provide an understanding of spatial distributions for evaluating monitoring data.     

Phytoplankton structure and dynamics in Lake Sanabria and Valparaíso reservoir (NW Spain)

The aim of this work is to compare the composition and seasonality of the phytoplankton population in a natural oligotrophic lake (Lake Sanabria) and a mesotrophic reservoir (Valparaíso). Both ecosystems are located on the Tera river course (NW Spain), which runs along an area of ancient metamorphic and plutonic rocks. Some physical and chemical parameters, chlorophyll a and phytoplankton biovolume were studied from monthly samples collected at different depths during the periods 1987–1989 (Lake Sanabria) and 1991–1992 (Valparaíso). Phytoplankton biovolume and chlorophyll a concentration were about five times higher in Valparaíso than in Lake Sanabria. Species composition (and main phytoplankton groups) were different. Valparaíso was highly dominated by diatoms and Lake Sanabria by cryptophytes and small chlorophytes. In spite of the fact that both sites were nitrogen limited, heterocystous cyanophytes (Anabaena sp.) were detected only in Valparaíso. The relationships between phytoplankton structure and trophic level, hydrological conditions and nitrate content are discussed.     

A chemotaxonomic method to quantify phytoplankton groups in freshwater lentic mesocosms: an approach including chlorophyll a breakdown products

The use of HPLC methods to determine and quantify phytoplankton population composition, is sometimes less time-consuming than microscopic identification. However, its general application poses problems since high discrepancies between chlorophyll a calculated using chemotaxonomic methods and direct measurements were noticed. For instance, chemotaxonomic protocols generally employed can lead to a poor estimation of total and relative abundance when high amounts of chlorophyll a breakdown products are present. Therefore, we propose a new approach to calculate relative abundance of algal groups in a phytoplankton population, based on integration of these degradation products in the chemotaxonomic assessment in lentic and shallow freshwater ecosystems.     

Nutrient addition effects on chlorophyll a,phytoplankton biomass,and heterocyte formation in Lake Erie’s central basin during 2014–2017: Insights into diazotrophic blooms in high nitrogen water

1. Phosphorus (P) usually is the primary limiting nutrient of phytoplankton biomass, but attention towards nitrogen (N) and trace nutrients, such as iron (Fe), has surfaced. Additionally, N-fixing cyanobacterial blooms have been documented to occur in N-rich, P-poor waters, which is counterintuitive from the paradigm that low N and high P promotes blooms. For example, Lake Erie's central basin has Dolichospermum blooms when nitrate concentrations are high, which raises questions about which nutrient(s) are selecting for Dolichospermum over other phytoplankton and why an N-fixer is present in high N waters?
2. We conducted a 4-year (2014–2017) study in Lake Erie's central basin to determine which nutrient (P, N, or trace nutrients such as Fe, molybdenum [Mo], and boron [B]) constrained chlorophyll concentration, phytoplankton biovolume, and nitrate assimilation using nutrient enrichment bioassays. The enriched lake water was incubated in 1-L bottles in a growth chamber programmed at light and temperatures of in situ conditions for 4–7 days. We also quantified heterocytes when N-fixing cyanobacteria were present.
3. Compared to the non-enriched control, the P-enriched (+P) treatment had significantly higher chlorophyll and phytoplankton biovolume in c. 75% of experiments. Combination enrichments of P with ammonium-N, nitrate-N, Fe, Mo, and B were compared to the +P treatment to determine secondary limitations. +P and ammonium-N and +P nitrate-N resulted in higher chlorophyll in 50% of experiments but higher phytoplankton biovolume in only 25% of experiments. These results show that P was the primary limiting nutrient, but there were times when N was secondarily limiting.
4. Chlorophyll concentration indicated N secondary limitation in half of the experiments, but biovolume indicated only N secondary limitation in 25% of the experiments. To make robust conclusions from nutrient enrichment bioassays, both chlorophyll and phytoplankton biovolume should be measured.
5. The secondary effects of Fe, Mo, and B on chlorophyll were low (<26% of experiments), and no secondary effects were observed on phytoplankton biovolume and nitrate assimilation. However, +P and Fe resulted in more chlorophyll than +P in experiments conducted during Dolichospermum blooms, and +P and B significantly increased the number of heterocytes in Dolichospermum. These results indicate that low Fe availability might select for Dolichospermum, and low B constrains heterocyte formation in the central basin of Lake Erie. Furthermore, these results could apply to other lakes with high N and low P where diazotrophic cyanobacterial blooms occur.

     

Responses of phytoplankton to fish predation and nutrient loading in shallow lakes: a pan-European mesocosm experiment

1. The impacts of nutrients (phosphorus and nitrogen) and planktivorous fish on phytoplankton composition and biomass were studied in six shallow, macrophyte‐dominated lakes across Europe using mesocosm experiments. 2. Phytoplankton biomass was more influenced by nutrients than by densities of planktivorous fish. Nutrient addition resulted in increased algal biomass at all locations. In some experiments, a decrease was noted at the highest nutrient loadings, corresponding to added concentrations of 1 mg L^?1 P and 10 mg L^?1 N. 3. Chlorophyll a was a more precise parameter to quantify phytoplankton biomass than algal biovolume, with lower within‐treatment variability. 4. Higher densities of planktivorous fish shifted phytoplankton composition toward smaller algae (GALD < 50 μm). High nutrient loadings selected in favour of chlorophytes and cyanobacteria, while biovolumes of diatoms and dinophytes decreased. High temperatures also may increase the contribution of cyanobacteria to total phytoplankton biovolume in shallow lakes.     

Responses to fish predation and nutrients by plankton at different levels of taxonomic resolution

1. To improve mechanistic understanding of plankton responses to eutrophication, a mesocosm experiment was performed in the shallow littoral zone of a south Swedish lake, in which nutrient and fish gradients were crossed in a fully factorial design. 2. Food chain theory accurately predicted total biomass development of both phyto‐ and zooplankton. However, separating zooplankton and algae into finer taxonomic groups revealed a variety of responses to both nutrient and fish gradients. 3. That both nutrients and fish are important for phytoplankton dynamics was seen more clearly when viewing each algal group separately, than drawing conclusions only from broad system variables such as chlorophyll a concentration or total phytoplankton biovolume. 4. In some taxa, physiological constraints (e.g. sensitivity to high pH and low concentrations of free CO₂) and differences in competitive ability may be more important for the biomass development than fish predation, grazing by herbivorous zooplankton, and nutrient availability. 5. We conclude that food chain theory accurately predicted responses in system variables, such as total zooplankton or algal biomass, which are shaped by the dynamics of certain strong interactors (‘keystone species’), such as large cladocerans, cyanobacteria and edible algae (<50 μm), whereas responses at finer taxonomic levels cannot be predicted from current theory.     

Classification and ordination of profundal macroinvertebrate communities in nutrient poor, oligo-mesohumic lakes in relation to environmental data

SUMMARY. 1. Profundal macroinvertebrates and water chemistry of sixty-eight nutrient poor, poorly buffered lakes were sampled in 1983 or 1986. Assemblages of profundal zoobenthos were classified using two-way indicator species analysis (TWINSPAN), ordinated by canonical correspondence analysis (CCA), and related to physico-chemical factors using CCA. discriminant analysis, and regression.
2. 90% of zoobenthos (TWINSPAN) groups were classified correctly using discriminant analysis. Depth (Function 1), soluble reactive Si(OH)₄ and HCO₃, -(2), and phytoplankton biovolume and pH (3) were the strongest correlates with the three discriminant functions.
3. Regression analysis showed that depth, KmnO₄, consumption, and phytoplankton biovolume were the best estimators of zoobenthos biomass in the profundal.
4. Multivariate analyses showed species assemblages amongst the profundal zoobenthos to be good indicators of lake type, particularly depth, pH, and phytoplankton biovolume.