Time-series measurement of grazing rates of zooplankton and bivalves

The current state of the art in automated measurement of grazingrates of zooplankton and bivalves is evaluated. Limitationsto the development of automated methods are discussed with referenceto theoretical considerations. Different approaches to time-seriesmeasurements are examined, and some inadequately investigatedtime scales for measurement of grazing rates are outlined. Levelsof automation of existing experimental systems are described,and a recently developed grazing system based on in vivo fluorescenceadvanced. The new system is suited to use with zooplankton orbivalves, but depends on precise calibration of in vivo fluorescence.It facilitates rapid measurement of functional response to foodconcentration, estimation of time-series rates at constant foodconcentrations, and measurement of rates while simulating patchyfood distribution. Results derived from experiments with krill(Nyctiphanes australis) and cockles (Chione stutchburyi) usingthis grazing system are presented, and some future directionsfor instrument development suggested.     

Type-3 functional response in limnetic suspension-feeders,as demonstrated by in situ grazing rates

Field-measured grazing rates (ml/animal/d) of cladocerans (mostly daphniids) and diaptomids were assembled from various published studies and plotted as a function of corresponding phytoplankton concentration (μg l⁻¹ f.w.). Filtering rates of both zooplankton groups initially increased with seston concentration until maximal grazing rates were observed at approximately 4 × 10² and 1 × 10² μg l⁻¹ for cladocerans and copepods, respectively; at higher algal concentrations, filtering rates of both declined as a function of food concentration. The shape of these curves are most consistent with Holling's (1966) Type 3 functional response. We found little support for the Type 3 functional response in published laboratory studies of Daphnia; most investigators report either a Type 1 or Type 2 response. The one study in which the Type 3 response was observed involved experiments where animals were acclimated at low food concentrations for 24 h, whereas those studies associated with response Types 1 or 2 had acclimation periods of only 1 to 3 h. We therefore assembled relevant data from the literature to examine the effect of acclimation period on the feeding rates of Daphnia at low food concentrations. In the absence of any acclimation, animals filtered at extremely low rates. After 2 h of acclimation, however, filtering rates increased 4 to 5-fold but declined again with longer durations; after > 70 h of pre-conditioning, filtering rates were almost as low as they had been with no acclimation. We also found little support for the Type 3 functional response in published studies of copepods. The only study associated with a Type 3 response involved a marine copepod that had been subjected to a starvation period of 48 h; however, an analysis of the effects of acclimation period did not yield conclusive evidence that filtering rates of freshwater copepods (Diaptomus and Eudiaptomus) decrease significantly with acclimation duration. The low filtering rates associated with long acclimation periods in laboratory experiments appears to be a direct result of animals becoming emaciated from prolonged exposure to low food concentrations, a situation which renders them incapable of high filtering rates. This may explain the Type 3 functional response for field cladocerans, since zooplankton in food-limiting situations are constantly exposed to low food concentrations, and would therefore have low body carbon and consequently less energy to filter-feed. We cannot, however, use this to explain the Type 3 response for field diaptomids, since copepods in the laboratory did not appear to lose body carbon even after 72 h of feeding at very low food levels, and there was inconclusive evidence that either Diaptomus or Eudiaptomus decrease their filtering rates with acclimation period. Although Incipient Limiting Concentrations (ILC) for Daphnia ranged from 1 to 8.5 × 10³ μg 1⁻¹, more than half of these fell between 1 and 3 × 10³ μg l⁻¹, bracketing the value of 2.7 × 10² μg l⁻¹ for field cladocerans. There was, however, a great deal of variation in reported maximum ingestion rates (MIR), maximum filtering rates (MFR) and ILC values for Daphnia magna. ILC values from the few laboratory studies of freshwater copepods ranged between 0.5 to 2.8 × 10³ μg 1⁻¹, and was higher than the ILC value of approximately 0.2 × 10³ μg l⁻¹ calculated for field populations of D. minutus. Generally, there was considerable agreement among laboratory studies regarding the shape of grazing-rate and ingestion-rate curves when data were converted to similar units and presented on standardized scales.     

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.     

Effects of UV-B irradiated algae on zooplankton grazing

We tested the effects of UV-B stressed algae on grazing rates of zooplankton. Four algal species (Chlamydomonas reinhardtii, Cryptomonas sp., Scenedesmus obliquus and Microcystis aeruginosa) were used as food and fed to three zooplankton species (Daphnia galeata, Bosmina longirostris and Brachionus calyciflorus), representing different taxonomic groups. The phytoplankton species were cultured under PAR conditions, and under PAR supplemented with UV-B radiation at two intensities (0.3 W m^–2 and 0.7 W m^–2, 6 hours per day). Ingestion and incorporation experiments were performed at two food levels (0.1 and 1.0 mg C l^–1) using radiotracer techniques. The effect of food concentration on ingestion and incorporation rate was significant for all three zooplankton species, but the effect of UV-B radiation was more complex. The reactions of the zooplankton species to UV-B stressed algae were different. UV-B stressed algae did not affect Daphnia grazing rates. For Bosmina the rates increased when feeding on UV-B stressed Microcystis and decreased when feeding on UV-B stressed Chlamydomonas, compared with non-stressed algae. Brachionus grazing rates were increased when feeding on UV-B stressed Cryptomonas and UV-B stressed Scenedesmus, and decreased when feeding on UV-B stressed Microcystis, compared with non-stressed algae. These results suggest that on a short time scale UV-B radiation may result in increased grazing rates of zooplankton, but also in decreased grazing rates. Long term effects of UV-B radiation on phytoplankton and zooplankton communities are therefore difficult to predict.     

Further observations on limnetic zooplankton in a small lake and two enclosures containing fish

SUMMARY. In a small lake (Blelham Tarn, English Lake District) two plastic cylinders, each of which extends from the lake surface to the bottom sediments and encloses 18 000 m³ of water, were used to investigate effects of enclosure on the zooplankton. This paper describes observations from March 1973 to October 1974 and is the second part of a report on a project spanning 4 years.
Seasonal population densities of the more abundant species were essentially similar to those observed in the first part of the project, despite large changes in algal and predator populations. In the enclosures, fish (mostly Perca fluviatilis ) populations are known from observations by divers to have been large in the period under consideration and small in the first part of the project. Populations of Chaoborus larvae in the enclosures were small at all seasons from March 1973 to October 1974.     

The dynamics and role of limnetic zooplankton in Loosdrecht lakes (The Netherlands)

The paper summarizes the results of a ten-year (1981–1991) zooplankton research on the Lake Loosdrecht, a highly eutrophic lake. The main cause of the lake's eutrophication and deteriorating water quality was supply up to mid 1984 of water from the River Vecht. This supply was replaced by dephosphorized water from the Amsterdam-Rhine Canal in 1984. The effects of this and other restoration measures on the lake's ecosystem were studied. Despite a reduction in the external P-load from ca. 1.0 g P m^–2 y^–1 to ca. 0.35 g m^–2 y^–1 now, the filamentous prokaryotes, including cyanobacteria and Prochlorothrix, continue to dominate the phytoplankton.Among the crustacean plankton Bosmina spp, Chydorus sp. and three species of cyclopoid copepods and their nauplii are quite common. Though there was no major change in the composition of abundant species, Daphnia cucullata, which is the only daphnid in these lakes, became virtually extinct since 1989. Among about 20 genera and 40 species of rotifers the important ones are: Anuraeopsis fissa, Keratella cochlearis, Filinia longiseta and Polyarthra. The rotifers usually peak in mid-summer following the crustacean peak in spring. The mean annual densities of crustaceans decreased during 1988–1991. Whereas seston (< 150 µm) mean mass in the lake increased since 1983 by 20–60%, zooplankton (> 150 µm) mass decreased by 15–35%.The grazing by crustacean community, which was attributable mainly to Bosmina, had mean rates between 10 and 25% d^–1. Between 42 and 47% of the food ingested was assimilated. In spring and early summer when both rotifers and crustaceans have their maximal densities the clearance rates of the rotifers were much higher. Based on C/P ratios, the zooplankton (> 150 µm) mass contained 2.5 times more phosphorus than seston (< 150 µm) mass so that the zooplankton comprised 12.5 % of the total-P in total particulate matter in the open water, compared with only 4.5% of the total particulate C. The mean excretion rates of P by zooplankton varied narrowly between 1.5 and 1.8 µg P 1^– d^–1, which equalled between 14 and 28% d^–1 of the P needed for phytoplankton production.The lack of response to restoration measures cannot be ascribed to one single factor. Apparently, the external P-loading is still not low enough and internal P-loading, though low, may be still high enough to sustain high seston levels. Intensive predation by bream is perhaps more important than food quality (high concentrations of filamentous cyanobacteria) in depressing the development of large-bodied zooplankton grazers, e.g. Daphnia. This may also contribute to resistance of the lake's ecosystem to respond to rehabilitation measures.     

The characterization of phosphorus excretion products of a natural population of limnetic zooplankton

The characteristics of phosphorus excretions of zooplankton collected in spring 1974 from Stonehouse Pond, New Hampshire were studied with gel filtration, ultraviolet spectroscopy and seston incorporation. The soluble phosphorus component released from a natural population has a molecular weight the same as orthophosphate and behaves similar to orthophosphate in seston incorporation studies. Approximately 15 percent of the total phosphorus released was organic, but could not be identified by ultraviolet spectroscopy. No evidence of nucleic acid excretion or any hydrolytic degradation product was detected in the ultraviolet spectrum.The experimental work was based on a thesis submitted to the Graduate School of the University of New Hampshire in partial fulfillment of the requirements for the degree of Doctor of Philosophy.     

Summer zooplankton dynamics in the limnetic and littoral zones of a humic acid lake

The limnetic and littoral zooplankton were studied during summer 1989 in Triangle Lake, a humic acid (pH 4.9) bog lake in Ohio, USA. The limnetic zooplankton showed low species richness and biomass, and dominance by the rotifers Kellicottia bostoniensis and Polyarthra vulgaris. In the littoral, species richness and biomass were markedly higher, and the crustaceans Alona guttata, Ceriodaphnia quadrangula, Chydorus sphaericus, Simocephalus serrulatus and Tropocyclops prasinus, and the rotifer Ascomorpha ecaudis were the dominants. The extreme rotifer dominance and lack of cladocerans in the limnetic zone were likely the result of Chaoborus predation. A pronounced mid-summer decline in cladoceran biomass in the littoral was likely due to predation by T. prasinus and Utricularia (bladderwort).     

Nonmetric multidimensional scaling analyses of temporal variation in the structure of limnetic zooplankton communities

Nonmetric multidimensional scaling analysis was used to graphically summarize and analyze seasonal changes in the structure of limnetic zooplankton communities in Blelham Tarn, English Lake District and two large experimental enclosures (tubes A and B) therein. Species abundances in weekly samples taken from June–December, 1976 from the three sites were ordinated. Paths through species space describing temporal changes in community structure indicated that the tube B community oscillated around some average composition and was dominated by small zooplankters whereas those from tube A and the tarn changed quite distinctly through the season and had higher predator densities. Examination of other characteristics of the sites indicated that the observed differences in seasonal cycles could be related to variations in the intensity of predation, the quality of food for herbivorous zooplankters and the dynamics of nutrient input. The importance of considering time variation in lake properties, particularly in assessing lake responses to stress and in typing lakes for experimentation or management, is emphasized.     

Comparison of carbon-specific growth rates and rates of cellular increase of phytoplankton in large limnetic enclosures

Potential carbon-specific growth rates of phytoplankton wereestimated from a series of measurements of photosynthetic radio-carbonuptake over 4- and 24-h exposure periods in the light fieldsof three large limnetic enclosures (‘Lund Tubes’),each providing different limnological and trophic conditions.Photosynthetic behaviour and short-term, chlorophyll-specificcarbon-fixation rates conformed to well-established criteriabut, over 24 h, the net retention represented 23–82% ofthe carbon fixed during the daylight hours. Potential mean growthrates (k'_p, of the photo-autotrophic community were calculatedas the net exponential rates of daily carbon-accumulation relativeto derived, instantaneous estimates of the cell carbon-content.Apparent actual community growth rates (k'_D were calculatedas the sum of the exponential rates of change of each of themajor species present, corrected for probable rates of in situgrazing and sinking, and expressed relative to the fractionof total biomass for which they accounted. The correspondingvalues were only occasionally similar, k'_p generally exceedingK'_D by a factor of between 1 and 30 or 1 and 14, depending uponthe carbon:chlorophyll ratio used. The ratio, K'_p/K'_D was foundto vary inversely both to k'_D and to k_n, the net rate of changein phytoplankton biomass, suggesting that measured carbon fixationrates merely represent a capacity for cellular increase which,owing to other likely limitations upon growth, is seldom realized.Apparent rates of loss of whole cells do not account for theloss of carbon; that the ‘unaccounted’ loss rates(K'_p–K'_D varied in direct proportion to K'_p (i.e., losseswere least when chlorophyll-specific photosynthetic productivitywas itself limited) is best explained by physiological voidingof excess carbon (for instance, by respiration, photorespiration,excretion) prior to the formation of new cells.     

Natural filtration rates of zooplankton in a closed system: the derivation of a community grazing index

Measurements of individual filtration rates by natural suspensionsof zooplankton are presented. The products filtration rate xanimal concentration, calculated for each of several species-sizecategories, are summated to estimate the total volume of waterfiltered by the zooplankton per unit time, here defined as the"Community Grazing Index" (CGI). Seasonally, CGI varies overnearly three orders of magnitude. The Daphnia hyalina individualspresent frequently contributed >90% of CGI. Individual filtrationrates were highest when the concentrations of suitable foodswere limiting (equivalent to 0.1–0.2 µg C ml^–1),but were depressed after long (3-week) periods of low food availabilityand when large Microcystis colonies dominated the phytoplankton.Approximately one order of magnitude separated these extremes.Fluctuations in CGI owe relatively more to change in the concentration,size and species distribution of the animal population thanto changes in individual filtration rate. ¹Present address: Anglian Water Authority, Oundle, PeterboroughPE8 4AS, UK     

Fixation artifacts of phytoplankton in zooplankton grazing experiments

The influence of various fixatives on size distribution and cell concentration was investigated using two types of electronic particle counters. Both in autotrophic and heterotrophic flagellates drastic losses of cells and changes of size were found in all preservatives used. Also preserved natural seston samples were not representative of the original fresh sample, and they continued changing and varying in the course of time. In diatoms also changes in size were noticed due to the fixative, however, doubt arises whether they were real or only measured differently by the particle counter. Even filtered sea water could not be preserved without obvious changes in particle distribution. Apart from diatoms, commonly used fixatives must be regarded worthless to preserve most phytoplankton and protozooplankton species in grazing experiments with zooplankton.     

Some studies on the ecology of limnetic crustacean zooplankton in Lake Begnas and Rupa,Pokhara Valley,Nepal

Monthly zooplankton samples taken during the period February, 1977 to February, 1978 in the deepest portion in two shallow sub-tropical Lakes, Begnas and Rupa in the Pokhara Valley, Nepal were studied. Four peaks of zooplankton abundance were noted. Adult and copepodid Copepoda were numerically dominant in both lakes with 56% Copepoda, 24% Cladocera and 20% nauplii in Lake Begnas and 48% Copepoda, 36% Cladocera and 16% nauplii in Lake Rupa. Other forms like Chaoborus larvae occurred sporadically in both lakes. An occurrence of the rare Limnocnida nepalensis (Coelenterata: Limnomedusae) in Lake Rupa was also noted during April and May, 1977. Although both of these lakes had already been ranked as eutrophic, the absence of calanoids, relative abundance of Bosmina longirostris and higher gross primary production in Lake Rupa is an indication of a higher trophic condition than that of Lake Begnas.     

浮游动物摄食在赤潮生消过程中的作用

浮游动物摄食在赤潮生消过程中起相当重要的作用。由于摄食过程的复杂性和生物物种与个体行为存在的多样性 ,使得赤潮过程中浮游动物的摄食研究具有相当难度。从浮游动物的摄食类型和习性、浮游动物摄食率测定、浮游动物选择性摄食对赤潮群落演替发展方向、浮游动物摄食在有毒微藻赤潮中的作用、浮游动物摄食在中国赤潮研究中的关键科学问题等几个方面探讨了浮游动物摄食对赤潮生物种群动力学的影响 ,为理解和治理赤潮提供科学依据     

乐清湾浮游动物的季节变动及摄食率

2002年8月、11月、2003年2月和5月,在乐清湾进行了4个航次生物、化学和水文等专业综合调查。根据采集的浮游动物样品的分析鉴定及海上现场实验结果,对浮游动物的群落组成、生物量、丰度、多样性指数的分布和季节变动及其浮游动物对浮游植物的摄食率进行研究。结果表明,乐清湾已鉴定的浮游动物有56属,75种,17类浮游幼体,主要可划分为4个生态类群,以近岸低盐类群为主,其优势种为真刺唇角水蚤Labidoceraeuchaeta、太平洋纺锤水蚤Acartiapacifica、驼背隆哲水蚤Acrocalanusgibber、中华假磷虾Pseudeuphausiasinica和百陶箭虫Sagittabedoti等,半咸水河口类群、暖水性外海种和广布种的种数相对较少。浮游动物生物量和丰度的平面分布趋势基本一致,有明显季节变化。2月份和5月份,浮游动物生物量和丰度,从湾顶向湾口呈逐渐增加趋势;8月份,生物量和丰度的分布与2月份、5月份的分布趋势不同,从湾顶向湾口,生物量和丰度逐渐降低;11月份,生物量和丰度的平面分布相对均匀。浮游动物种类多样性指数有明显的季节变化,其动态变化与浮游动物种数和丰度的变化一致。微型浮游动物对浮游植物存在摄食压力,且摄食率有季节变化,摄食率的变化在0.15~0.48d-1。     

Diel patterns of zooplankton grazing in a shallow lake

A diel survey of in situ species-specific zooplankton clearancerates (with radioactively labelled Chlorella cells) was conductedin the shallow lake of Cr?teil characterized by small-sizedplanktonic forms (algae<10µm and zooplankton <1.3mm). Experiments were performed every 4h at two depths (1 and4m). Power functions relating individual filtering rates tobody length were established for the three most abundant cladoceransand for calanoids (nauplii being included in this feeding group),for each depth and time. No filtering penodicity was observedin Ceriodaphnia spp., adults and copepodites of Eudiaptomusgracilis and Eurytemora velox, and nauplii. On the contrary,clear nocturnal filtering peaks were obtained for Daphnia spp.and for Diaphanosoma brachyurum, these being more pronouncedfor the larger individuals at 1 m deep. The observed diel periodicitycannot be explained by variations in physico-chemical parametersor food concentration alone. The ecological significance ofthis phenomenon in the polymictic lake of Cr?teil is discussedin the light of previously published data and the hypothesesaccounting for it.     

浮游动物诱发藻类群体的形成

从研究蓝藻水华形成机理的需要出发,综述了浮游动物的牧食压力对藻类群体形成的诱发作用。指出诱发藻类群体形成的化合物来自牧食性浮游动物对藻类的有效牧食,是藻类群体形成的重要原因之一,而这些诱发性的化合物并不是有关生物体的组成成分,是种间相互作用的结果。藻类群体的形成方式有源于一个母细胞的分裂和业已存在的单细胞的聚合两种方式,栅藻的诱发性群体可能是来自一个母细胞的分裂,而在其它藻类的诱发性群体形成如铜绿微囊藻则可能是业已存在的单细胞的聚合。由于藻类形成群体后能显著降低浮游动物对其牧食速率,因此,这种诱发性群体形成的现象,可以解释为藻类对变化的牧食压力的一种有效的反牧食防御策略,也是两者协同进化的结果。浮游动物对藻类群体形成的重要作用,在研究模拟蓝藻群体及水华形成值得借鉴应用。作者还提出推测,水华蓝藻的群体形成,可能就是在富营养化条件下藻类快速生长,加上浮游动物的牧食压力共同作用下联合驱动的结果,而这种群体形成很可能在积累到一定程度后,结合特定的气象水文等理化因子,就会聚集于水表“爆发”出肉眼可见的水华。因此,开展浮游动物牧食作用对水华蓝藻早期群体形成诱发效应的研究不仅能加深对水华形成的全面认识,而且对于进一步认识藻类的诱发性反牧食防御适应机制、揭示生态系统中生物之间的复杂关系也具有重要的理论意义。     

A field technique for the determination of zooplankton grazing on natural bacterioplankton

SUMMARY. 1. A method for the determination of zooplankton community grazing rates on natural populations of bacteria is described.
2. Undisturbed zooplankton samples were incubated with ³H-thymidine-labelled bac teriioplankton, and animals and their food subsequently separated by size.
3. Prehandling of zooplankton by concentration before incubation reduce the grazing rates by a factor ranging from three to four.
4. Time series experiments showed that appropriate incubation periods ranged from 40 min at 14°C to 60 min at 10°C.
5. Spatial variability in the zooplankton was the main source of the overall standard error. This was considerably reduced by using composite samples.     

FO-spectra of chlorophyll fluorescence for the determination of zooplankton grazing

In the PHYTO-PAM phytoplankton analyzer the minimal fluorescence of dark-adapted samples (F₀) was assessed, which gives direct information on the chlorophyll-a content. Clearance rates (CR) of Daphnia and Brachionus were calculated from a decrease in chlorophyll-a concentration using the PHYTO-PAM fluorometer for non-sacrificial sampling of chlorophyll-a. Clearance rates of Daphnia were measured and compared with those based on the cell-counts method using an electronic particle counter (Coulter counter). Chlorophyll fluorescence-based CR for Daphnia magna were very strongly correlated with Coulter-based CR, signifying the potential suitability of the PHYTO-PAM in grazing experiments. A procedure for determination of rotifer clearance rates was developed and the effects of rotifer density, duration of the grazing period, and food concentration on CR were investigated. Between 10 and 30 rotifers in 2.5 ml food suspension (i.e. 4–12 rotifers per ml) appeared optimal for calculating CR. The application of the deconvolution of F₀-spectra in food selectivity experiments was evaluated using various mixtures of the green alga Scenedesmus obliquus and the cyanobacterium Microcystis aeruginosa fed to Brachionus. CR for Brachionus on M. aeruginosawere lower than on S. obliquusbut this was not caused by toxicity, because no mortality was observed. The higher CR on Scenedesmus than on Microcystis in the mixtures suggested selectivity. The importance of digital suppression of background fluorescence is highlighted in additional experiments with Daphnia feeding on mixtures of Microcystis and Scenedesmus, or on Microcystis alone. Without background correction of filtered samples, negative clearance rates were obtained for the `blue' Microcystis signal. Soluble fluorescing compounds of cyanobacterial origin, phycocyanin, were released from the Daphniaand contributed 40% to the overall-fluorescence. Deconvolution of F₀-spectra for the determination of chlorophyll-a using the PHYTO-PAM appears to be a suitable tool for determination of rotifer CR even at very low food concentrations. A drawback of the method is that rather high rotifer densities are required. The required grazing period, however, is shorter than for cell-count methods, the method is sensitive, clearance rates can be measured at low food concentrations (< 0.1 mg C l^–1) and information on selective feeding can be obtained.     

Calculations of zooplankton grazing rates according to a closed,steady-state,three-compartment model applied to different14C methods

A re-examination of the numerical example of the three-compartment model by CONOVER & FRANCIS (1973) showed that the warning by these authors for the misuse of radio isotopes in transfer studies within food chains is incorrect and based on a misinterpretation of their results. There is no difference in the estimate of transfer rate by use of specific activities or by use of total radioactivities observed in each compartment.After adapting the formulae developed by CONOVER & FRANCIS, their model was used to illustrate deviations of the programmed grazing rate in 3 types of grazing experiments; a) with¹⁴C present only in the phytoplankton at the start of the experiment, b) with¹⁴C only in the water, and c) with¹⁴C in both phytoplankton and water. Up to a duration of the grazing experiment of 2 hours, and at various light conditions and grazing pressures, deviations were small and did not exceed 4%. These results are not directly applicable to practical work because the quantitatively important loss by egestion of radioactive material was not accounted for, only losses by respiration were incorporated in the closed, steady-state model.Best calculations of the community filtering rate (fraction of the volume of the grazing vessel swept clear per day) were generally obtained with the formula (with t in hours) ((DPM zoo at time t)/(DPM phyto at 0+DPM phyto at t)/2)×24/t, applicable to all three types of grazing experiments considered.