Structure and functioning of the microbial loop in a boreal reservoir

The abundance and biomass of the main components of the microbial plankton food web (“microbial loop”)—heterotrophic bacteria, phototrophic picoplankton and nanoplankton, heterotrophic nanoflagellates, ciliates and viruses, production of phytoplankton and bacterioplankton, bacterivory of nanoflagellates, bacterial lysis by viruses, and the species composition of protists—have been determined in summer time in the Sheksna Reservoir (the Upper Volga basin). A total of 34 species of heterotrophic nanoflagellates from 15 taxa and 15 species of ciliates from 4 classes are identified. In different parts of the reservoir, the biomass of the microbial community varies from 26.2 to 64.3% (on average 45.5%) of the total plankton biomass. Heterotrophic bacteria are the main component of the microbial community, averaging 63.9% of the total microbial biomass. They are the second (after the phytoplankton) component of the plankton and contribute on average 28.6% to the plankton biomass. The high ratio of the production of heterotrophic bacteria to the production of phytoplankton indicates the important role of bacteria, which transfer carbon of allochthonous dissolved organic substances to a food web of the reservoir.     

Lysogeny and lytic viral production during a bloom of the cyanobacterium Synechococcus spp

Lytic viral production and lysogeny were investigated in cyanobacteria and heterotrophic bacteria during a bloom of Synechococcus spp. in a pristine fjord in British Columbia, Canada. Triplicate seawater samples were incubated with and without mitomycin C and the abundances of heterotrophic bacteria, cyanobacteria, total viruses and infectious cyanophage were followed over 24 h. Addition of mitomycin C led to increases in total viral abundance as well as the abundance of cyanophages infecting Synechococcus strain DC2. Given typical estimates of burst size, these increases were consistent with 80% of the heterotrophic bacteria and 0.6% of Synechococcus cells being inducible by the addition of mitomycin C. This is the highest percentage of lysogens reported for a natural microbial community and demonstrates induction in a marine Synechococcus population. It is likely that the cyanophage production following the addition of mitomycin C was much higher than that titered against a single strain of Synechococcus; hence this estimate is a minimum. In untreated seawater samples, lytic viral production was estimated to remove ca. 27% of the gross heterotrophic bacterial production, and a minimum of 1.0% of the gross cyanobacterial production. Our results demonstrate very high levels of lysogeny in the heterotrophic bacterial community, outside of an oligotrophic environment, and the presence of inducible lysogens in Synechococcus spp. during a naturally occurring bloom. These data emphasize the need for further examination of the factors influencing lytic and lysogenic viral infection in natural microbial communities.     

Temporal stability of marine phytoplankton in a subtropical coastal environment

We investigated the temporal stability of phytoplankton at a subtropical coastal site for 9 months by conducting chlorophyll and flow cytometric measurements at relatively high frequency (roughly at 2–5 day interval). Phytoplankton cells were grouped based on their sizes obtained from flow cytometric signals. We also conducted dilution experiments to estimate the growth and grazing mortality rates of different phytoplankton groups to assess whether the temporal stability of phytoplankton abundances was related with phytoplankton growth/grazing rates. Based on size-fractionated chlorophyll measurements, there was some indication that smaller phytoplankton cells were more stable than larger ones. However, by cytometric counting, there was no evidence for greater stability in small cells. Synechococcus, which had the lowest stability and dominated the <1 μm size class, showed a strong seasonal cycle that was highly dependent on temperature whereas eukaryotes did not have evident seasonal cycles. In general, biomass of a phytoplankton group consisting of several size classes was more stable than that of its sub component, consistent with the hypothesis that higher diversity leads to higher stability, probably related with the effect of statistical averaging (portfolio effect). Stability of heterotrophic bacteria was much higher than that of phytoplankton, leading to the speculation that bacteria were more diverse than phytoplankton. Phytoplankton stability was not related with their growth or grazing mortality rates. Our study suggests that species diversity should be taken into account when considering the temporal stability of phytoplankton.     

Analysis of the importance of spatial and temporal heterogeneity in the estimation of annual production by phytoplankton in a small,enriched, marine basin

In the Bedford Basin, Nova Scotia — a small, enriched, marine inlet — the annual production of phytoplankton was 220 g Cm⁻²yr⁻¹ with a standard deviation of ± 35 gC m⁻² yr⁻¹. The relative contributions of spatial and temporal fluctuations to the variance of the estimate of annual production are assessed, and conclusions are drawn on the design of sampling programs. Although on any given day differences were observed in production rate among four stations, the estimates of annual production agreed within 5 %. The results from Bedford Basin are compared with those from neighbouring St Margaret's Bay, which is relatively unenriched.     

The seasonal sensitivity of Cyanobacteria and other phytoplankton to changes in flushing rate and water temperature

The phytoplankton lake community model PROTECH (Phytoplankton RespOnses To Environmental CHange) was applied to the eutrophic lake, Esthwaite Water (United Kingdom). It was validated against monitoring data from 2003 and simulated well the seasonal pattern of total chlorophyll, diatom chlorophyll and Cyanobacteria chlorophyll with respective R²‐values calculated between observed and simulated of 0.68, 0.72 and 0.77 (all P<0.01). This simulation was then rerun through various combinations of factorized changes covering a range of half to double the flushing rate and from ?1 to +4 °C changes in water temperature. Their effect on the phytoplankton was measured as annual, spring, summer and autumn means of the total and species chlorophyll concentrations. In addition, Cyanobacteria mean percentage abundance (%Cb) and maximum percentage abundance (Max %Cb) was recorded, as were the number of days that Cyanobacteria chlorophyll concentration exceed two World Health Organization (WHO) derived risk thresholds (10 and 50 mg m^?3). The phytoplankton community was dominated in the year by three of the eight phytoplankton simulated. The vernal bloom of the diatom Asterionella showed little annual or seasonal response to the changing drivers but this was not the case for the two Cyanobacteria that also dominated, Anabaena and Aphanizomenon . These Cyanobacteria showed enhanced abundance, community dominance and increased duration above the highest WHO risk threshold with increasing water temperature and decreasing flushing rate: this effect was greatest in the summer period. However, the response was ultimately controlled by the availability of nutrients, particularly phosphorus and nitrogen, with occasional declines in the latter's concentration helping the dominance of these nitrogen‐fixing phytoplankton.     

Phytoplankton in the Marginal Ice Zone of the Greenland Sea during summer, 1984

Summary Phytoplankton biomass and species composition were studied in transects through the ice edge region of the Greeland Sea from 19 July to 8 August 1984. Biomass was estimated by vertical in situ chlorophyll fluorescence and pigment extraction of discrete samples. Preserved material was used for identification of phytoplankton species and calculation of their relative abundances. The results suggest that the various geographical regions of the Greenland Sea differ considerably in their phytoplankton development. Autotrophic biomass and species composition were closely associated with the extent of the annual and seasonal ice cover, hydrographic conditions, nutrient availability and the water masses typical of the different domains. In the NE Greenland polynya a deep mixed layer inhibited the development of a phytoplankton bloom, whereas greatest biomass concentrations were associated with a receding ice edge on the E Greenland Shelf. In the Fram Strait, the position of the relatively stationary ice edge is controlled by frontal dynamics, currents and wind. Due to rapidly changing physical and chemical conditions, phytoplankton biomass showed great variability between stations. High chlorophyll a concentrations may develop locally where melting ice causes stratification or can result from passive accumulation in eddies. In July/August 84 the Fram Strait area was dominated by a typical summer population of flagellates and large diatom species.Contribution 6 of the Alfred-Wegener-Institute for Polar and Marine Research     

Changes to the phytoplankton assemblage of Lake Kinneret after decades of a predictable, repetitive pattern

1. Phytoplankton abundance and species composition in Lake Kinneret, Israel, have been monitored at weekly or fortnightly intervals since 1969. This paper summarises the resulting 34‐year phytoplankton record with a focus on the last 13 years of new data, and reassesses an earlier conclusion that the lake phytoplankton shows remarkable stability despite a wide range of external pressures. 2. The Kinneret phytoplankton record can be split into two major periods. The first, from 1969 till 1993, was a period of distinct stability expressed by a typical annual pattern revolving around a spring bloom of the dinoflagellate Peridinium gatunense that repeated each year. The second period, starting around 1994 and ongoing, is characterised by the loss of the previously predictable annual pattern, with both ‘bloom years’ and ‘no‐bloom years’. 3. In the second period, deviations from the previous annual pattern include: the absence of the prevailing spring P. gatunense blooms in some years and increased variability in the magnitude of the bloom in others; intensification of winter Aulacoseira granulata blooms; higher summer phytoplankton biomass with replacement of mostly nanoplanktonic, palatable forms by less palatable forms; new appearance and establishment of toxin‐producing, nitrogen fixing cyanobacteria in summer; increase in the absolute biomass and percentage contribution of cyanobacteria to total biomass; and fungal epidemics attacking P. gatunense. 4. The 34‐year record serves to validate Schindler's (1987) assessment that phytoplankton species composition will respond to increased anthropogenic stress before bulk ecosystem parameters.     

Relationship between nutrient concentration,phytoplankton density,and zooplankton density in nutrient enriched experimental ponds

The effects of different levels of nutrient input on the plankton community was investigated in a two-year controlled fertilization study of eight experimental ponds. There were four treatments, each replicated: a control, to which no fertilizer was added, and three levels of nutrient addition. Limnological parameters including phytoplankton and zooplankton densities were measured frequently during both summers and less frequently during the rest of the year. Inorganic nitrogen and phosphorus concentrations in the treated ponds increased. Phytoplankton and zooplankton density increased with treatment level but was variable. There was a limited relationship between the average chlorophyll a concentration per summer and the average cladoceran dry weight per summer. Above chlorophyll a concentrations greater than 60–70 mg/m³ other factors such as a pH zooplankton mortality effect, prevailed.     

Virioplankton: Viruses in Aquatic Ecosystems

The discovery that viruses may be the most abundant organisms in natural waters, surpassing the number of bacteria by an order of magnitude, has inspired a resurgence of interest in viruses in the aquatic environment. Surprisingly little was known of the interaction of viruses and their hosts in nature. In the decade since the reports of extraordinarily large virus populations were published, enumeration of viruses in aquatic environments has demonstrated that the virioplankton are dynamic components of the plankton, changing dramatically in number with geographical location and season. The evidence to date suggests that virioplankton communities are composed principally of bacteriophages and, to a lesser extent, eukaryotic algal viruses. The influence of viral infection and lysis on bacterial and phytoplankton host communities was measurable after new methods were developed and prior knowledge of bacteriophage biology was incorporated into concepts of parasite and host community interactions. The new methods have yielded data showing that viral infection can have a significant impact on bacteria and unicellular algae populations and supporting the hypothesis that viruses play a significant role in microbial food webs. Besides predation limiting bacteria and phytoplankton populations, the specific nature of virus-host interaction raises the intriguing possibility that viral infection influences the structure and diversity of aquatic microbial communities. Novel applications of molecular genetic techniques have provided good evidence that viral infection can significantly influence the composition and diversity of aquatic microbial communities.     

南黄海潮汐锋对浮游细菌生物量分布的影响

2 0 0 1年 5月 16～ 2 3日、6月 10～ 2 4日和 2 0 0 2年 6月 5～ 12日 ,利用“北斗号”船只对南黄海鱼产卵场进行了 3次专项调查。研究了潮汐锋断面叶绿素 a浓度、浮游细菌生物量的分布 ,目的是阐明潮汐锋的存在对浮游细菌生物量分布的影响。3个航次中的叶绿素 a浓度变化范围分别是 0 .0 6～ 2 .34mg/ m3(2 0 0 1- 0 5 )、0 .0 8～ 0 .9mg/ m3(2 0 0 1- 0 6 )、0 .14～ 3.0 4 mg/ m3(2 0 0 2 - 0 6 )。 3航次的聚球藻 (Synechococcus spp.)蓝细菌生物量变化范围分别为 7.6 2～ 2 2 .0 6 mg C/ m3(2 0 0 1- 0 5 )、8.5 3～2 7.5 2 mg C/ m3(2 0 0 1- 0 6 )、0 .6 9～ 5 5 .90 m g C/ m3(2 0 0 1- 0 6 )。异养细菌生物量变化范围分别为 7.5 6～ 5 1.82 mg C/ m3(2 0 0 1-0 5 )、8.5 4～ 2 4 .77mg C/ m3(2 0 0 1- 0 6 )、3.12～ 10 .0 5 mg C/ m3(2 0 0 2 - 0 6 )。而聚球藻蓝细菌对浮游植物总生物量的贡献 (CB:PB)平均值分别为 :5 8% (2 0 0 1- 0 5 )、77% (2 0 0 1- 0 6 )、31% (2 0 0 2 - 0 6 )。结果表明 :南黄海鱼产卵场在春末夏初 (5～ 6月份 ) ,叶绿素 a浓度最大值及次大值主要分布在锋区及其邻近的层化区2 0 m以浅位置 ;聚球藻蓝细菌生物量最大值主要分布于锋区及层化区表层和水体中层 ;异养     

Nutrient-phytoplankton relationships in a tropical meromictic soda lake

Seasonal variation through one year in total nitrogen (TN), total phosphorus (TP), phytoplankton biomass, phytoplankton species composition and other environmental factors were examined in Lake Sonachi, a tropical meromictic soda lake. Mean concentrations of TN and TP were 11 000 µg N l^-1 and 100 µg P l^-1, respectively. Maximum concentrations of TN and TP occurred in the monimolimnion. Phytoplankton biomass ranged from 350 to 1260 mg m^-3. Synechococcus bacillaris, a small coccoid cyanophyte, dominated the phytoplankton. The mean chlorophyll a concentration of 37 mg · m^-3 was a modest value when compared with those of other tropical soda lakes. High TN:TP ratios indicated phosphorus limitation in the lake.     

Productivity of clear and humic lakes: nutrients,phytoplankton, bacteria

The relationships between long-term surface average concentrations of humic acids measured as water colour, dissolved organic carbon (DOC) or Secchi disk transparency and trophic state variables were studied with literature data from more than 600 freshwater lakes. The geometric means of summer surface average nutrient (phosphorus and nitrogen) concentration, phytoplankton biomass (chlorophyll concentration), and hypolimnetic anoxia (anoxic factor) were significantly higher in coloured than in clear lakes. The regressions of colour or DOC on these trophic state variables were positive and significant throughout a range of three orders of magnitude. Phytoplankton or primary productivity was higher in coloured lakes, when expressed per volume of epilimnion. Annual integral primary productivity expressed on an areal basis was smaller in coloured lakes, probably a reflection of shallower phototrophic depths in these lakes. There is evidence that annual integral bacteria productivity is much higher in coloured lakes for two reasons: first, epilimnetic bacteria production was ca. four times higher in coloured lakes, second, other studies have shown that hypolimnetic bacteria production is commonly higher than epilimnetic production, especially in anoxic hypolimnia that are frequent in coloured lakes. All volumetrically expressed variables indicated higher productivity in coloured lakes. In addition, high bacteria productivity reflects a different food chain involving mixotrophs, possibly compensating for low light conditions. Our analyses indicate that primary and secondary productivity is as high as or higher than in clear lakes. This revised version was published online in July 2006 with corrections to the Cover Date.     

Phytoplankton community growth-rate response to nutrient pulses in a shallow turbid estuary, Galveston Bay, Texas

Phytoplankton growth is a physiological process often limitedby temperature, nutrients or light, while biomass accumulationis a function of growth rates, grazing and deposition. Althoughprimary productivity measurements are usually used to assessresponses to limiting factors, the rates are proportional tobiomass and inversely related to grazing pressure during experimentalincubations. Alternatively, carbon-specific growth-rate determinationsprovide insights into physiological responses without the confoundingeffects of biomass and grazing. The objective of this studywas to quantify the growth-rate responses of phytoplankton toenhanced nutrient availability (nitrate and phosphate) overa range of in situ irradiances. Growth rates were determinedbased on chlorophyll a-specific ¹⁴C-uptake rates by phytoplankton.Phytoplankton demonstrated high (24 h) growth rates when exposedto increased concentrations of limiting nutrients, independentof the surface irradiances (12–41%). Growth-rate responseswere also compared with the biomass (chlorophyll a) responsesand community composition. Observed and estimated phytoplanktonbiomass changes during the incubations differed, emphasizingthe structural role of grazers on the phytoplankton community.The phytoplankton community in Galveston Bay has the potentialto instantaneously respond to nutrient pulses, facilitatingdiatom biomass accumulations in spring and summer and small,flagellated species and cyanobacteria during periods of lownutrient inputs. Thus, Galveston Bay phytoplankton biomass andcommunity composition reflect a dynamic balance between thefrequency of nutrient pulsing and grazing intensity.     

Coherent assembly of phytoplankton communities in diverse temperate ocean ecosystems

The annual cycle of phytoplankton cell abundance is coherent across diverse ecosystems in the temperate North Atlantic Ocean. In Bedford Basin, on the Scotian Shelf and in the Labrador Sea, the numerical abundance of phytoplankton is low in spring and high in autumn, thus in phase with the temperature cycle. Temperature aligns abundance on a common basis, effectively adjusting apparent cell discrepancies in waters that are colder or warmer than the regional norm. As an example of holistic simplicity arising from underlying complexity, the variance in a community variable (total abundance) is explained by a single predictor (temperature) to the extent of 75% in the marginal seas. In the estuarine basin, weekly averages of phytoplankton and temperature computed from a 13 year time-series yield a predictive relationship with 91% explained variance. Temperature-directed assembly of individual phytoplankton cells to form communities is statistically robust, consistent with observed biomass changes, amenable to theoretical analysis, and a sentinel for long-term change. Since cell abundance is a community property in the same units for all marine microbes at any trophic level and at any phylogenetic position, it promises to integrate biological oceanography into general ecology and evolution.     

The significance of viruses to mortality in aquatic microbial communities

A variety of approaches including enumeration of visibly infected microbes, removal of viral particles, decay of viral infectivity, and measurements of viral production rates have been used to infer the impact of viruses on microbial mortality. The results are surprisingly consistent and suggest that, on average, about 20% of marine heterotrophic bacteria are infected by viruses and 10–20% of the bacterial community is lysed daily by viruses. The effect of viruses on phytoplankton is less certain, but ca. 3% of Synechococcus biomass may be lysed daily. The fraction of primary productivity this represents depends upon the relative biomass and growth rate of Synechococcus. Virus enrichment experiments suggest that the productivity of eukaryotic phytoplankton would be ca. 2% higher in the absence of viruses. Overall, probably about 2–3% of primary productivity is lost to viral lysis. There is considerable variation about these estimates; however, they represent a starting point for incorporating viral-mediated processes into aquatic ecosystem models.     

Spatial,temporal, and storm runoff-related variations in phytoplankton community structure in a small,suburban reservoir

Phytoplankton abundance and community structure were determined routinely over an annual period and intensively during two storm-runoff events in a small suburban reservoir in northern Virginia, U.S.A. Traditional graphical techniques and a multivariate approach (Principal Components Analysis) were used to demonstrate a seasonal pattern of phytoplankton succession with greens and blue-greens dominant in summer, diatoms and chrysophytes in spring and fall, and cryptophytes in winter. Spatial variations were minor over horizontal and vertical dimensions during spring mixis, but depth variations were substantial during summer stratification. Storm runoff had little effect on phytoplankton composition during the stable summer period, but was associated with a substantial perturbation in community structure during the spring to summer transition.     

Distribution Pattern of Photosynthetic Picoplankton and Heterotrophic Bacteria in the Northern South China Sea

The environmental regulation of plcoplankton distribution in the northern South China Sea was examined In winter and summer of 2004. The average abundance of Synechococcus, Prochlorococcus, and heterotrophlc bacteria was lower In winter （30, 21, and 780×10^3 cells/cm^3, respectively） than In summer （53, 85, and 1 090×10^3 cells/cm^3, respectively）, but the seasonal pattern was opposite for plcoeukaryotlc phytoplankton （4 500 and 3 200 cells/cm^3 In winter and summer, respectively）. Synechococcus, picoeukaryotes, and bacteria were most abundant in the nutrient-rich coastal zone and continental shelf, but Prochlorococcus was most abundant In the continental slope and open ocean. The vertical distribution of each photosynthetic group and heterotrophlc bacteria changed between the two seasons. Synechococcus populations with apparently different phycoerythrobilin content occurred at many stations In the summer. In addition, two different populations of Prochlorococcus were found： （i） small, weakly fluorescing cells in the surface layer; and （ii） larger, strongly fluorescent cells In the deep layer. The distribution pattern of photosynthetic plcoplankton and heterotrophlc bacteria depends on environmental effects and their ecophyslologlcal differences. The distribution of Synechococcus appeared to be related to nutrient availability, whereas the distribution of Prochlorococcus appeared to be limited by temperature. Synechococcus was the only plcophytoplankton with a consistent strong relationship with bacteria.     

Food Web Structure in the Recently Flooded Sep Reservoir as Inferred From Phytoplankton Population Dynamics and Living Microbial Biomass

Phytoplankton dynamics, bacterial standing stocks and living microbial biomass (derived from ATP measurements, 0.7-200 mm size class) were examined in 1996 in the newly flooded (1995) Sep Reservoir ('Massif Central,' France), for evidence of the importance of the microbial food web relative to the traditional food chain. Phosphate concentrations were low, N:P ratios were high, and phosphate losses converted into carbon accounted for <50% of phytoplankton biomass and production, indicating that P was limiting phytoplankton development during the study. The observed low availability of P contrasts with the high release of "directly" assimilable P often reported in newly flooded reservoirs, suggesting that factors determining nutrient dynamics in such ecosystems are complex. The phosphate availability, but also the water column stability, seemed to be among the major factors determining phytoplankton dynamics, as (i) large-size phytoplankton species were prominent during the period of increasing water column stability, whereas small-size species dominated phytoplankton assemblages during the period of decreasing stability, and (ii) a Dinobryon divergens bloom occurred during a period when inorganic P was undetectable, coinciding with the lowest values of bacterial standing stocks. Indication of grazing limitation of bacterial populations by the mixotrophic chrysophyte D. divergens (in late spring) and by other potential grazers (mainly rotifers in summer) seemed to be confirmed by the Model II or functional slopes of the bacterial vs phytoplankton regressions, which were always <0.63. Phytoplankton biomass was not correlated with phosphorus sources and its contribution was remarkably low relative to the living microbial biomass which, in contrast, was positively correlated with total phosphorus in summer. We conclude that planktonic microheterotrophs are strongly implicated in the phosphorus dynamics in the Sep Reservoir, and thus support the idea that an important amount of matter and energy flows through the "microbial loop" and food web, shortly after the flooding of a reservoir.     

Bioassay for phosphate demand in phytoplankton from acidified lakes: Lake Njupfatet,an example of phosphate deficiency induced by liming

A bioassay was developed, involving steady-state ATP level determinations, for estimation of phosphate demand and deficiency in natural phytoplankton communities. The studies were performed on phytoplankton from the moderately acidified Lake Njupfatet in central Sweden before and after liming. Phytoplankton samples from in situ enclosure experiments with low-dose enrichments of nitrate and phosphate and removal of large (> 100 µm) zooplankton and from the lake water were collected. The phytoplankton were concentrated by through-flow centrifugation and post-cultured in the laboratory with or without the addition of phosphate. A relative increase in the ATP:chlorophyll a ratio after the phosphate treatment as compared to samples without phosphate enrichment was found to be a highly reproducible indicator of phosphate deficiency in the natural phytoplankton population. In contrast, the absolute ATP:chlorophyll a ratio varied substantially between different sampling occasions. No phosphate deficiency was detected in phytoplankton from the acidic lake or from fertilized in situ enclosures. However, phytoplankton from in situ enclosures without added nutrients showed evidence of phosphate limitation after 21 days incubation. Also, the phytoplankton community developed a significant phosphate deficiency the summer after lake liming. The results from the ATP analyses are compared with chemical data of the lake water, phytoplankton community structure and phosphatase activities in the lake before and after liming. The average total biomass of phytoplankton and the average Tot-P measured during May to September decreased with appr. 30% after liming while Tot-N was essentially unaffected and the phosphatase activities increased by 1000–2000%.     

REACTIVE OXYGEN METABOLISM IN THE TROPICAL BROWN ALGA DICTYOTA DICHOTOMA

There is a growing understanding that phagotrophic ciliates are often important members of aquatic communities in terms of their trophic role and mobilization of small cell production to higher consumers. As formidable consumers of small phytoplankton species they are likely to be also important in determining the community composition of the pico- and nanophytoplankton assemblages. Dilution method experiments were conducted during the winter and summer in the South Slough, an arm of the Coos Bay on the southern Oregon coast, to assess the impact of ciliate grazing on two size fractions of chlorophyll (0.2 to 5 mm and> 5 mm) and on the growth and abundance of specific phytoplankton groups, particularly cryptophytes and Synechococcus sp. The premise of the dilution technique is that grazers are diluted with their food and the observed rate of change in chlorophyll or phytoplankton abundance is linearly related to the dilution factor. Results from previous studies using the dilution technique have been given in terms of the grazing impact of microzooplankton on total chlorophyll. The findings of the research presented using a more rigorous application of the dilution method suggest that ciliates are differential in their grazing of phytoplankton, targeting small phytoplankton biomass and preying selectively on components of the assemblage that constitute this biomass.