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.     

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 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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

Seasonal dynamics and grazing rate of zooplankton in Yueqing Bay, China

The species composition, biomass, abundance, and species diversity of zooplankton were determined for samples collected from August 2002 to May 2003 from 14 stations in Yueqing Bay, China. Phytoplankton growth rate and microzooplankton grazing rate were obtained by using the dilution method developed by Landry and Hassett. The spatial and temporal variations of zooplankton and its relationship with environmental factors were also analyzed. The results showed that the zooplankton in the Yueqing Bay could be divided into four ecotypes, namely coastal low saline species, estuary brackish water species, offshore warm water species, and eurytopic species. A total of 75 species of zooplankton belonging to 56 genera and 17 groups of pelagic larva were identified in the Yueqing Bay. The coastal low saline species was the dominant ecotype in the study area, and the dominant species were Labidocera euchaeta, Acartia pacifica, Acrocalanus gibber, Pseudeuphausia sinica, and Sagitta bedoti among others. There was considerable seasonal variation in zooplankton biomass and abundance in the surveyed areas. The peak biomass appeared in August, descending in November and in May, and the lowest biomass appeared in February. Similarly, the highest abundance of zooplankton was observed in August, with the abundance descending in the following months: May, November, and February. There were similar horizontal distribution patterns for the biomass and the abundance of zooplankton. They both increased from the upper to the lower bay in February and May, but decreased from the upper to the lower bay in August. Biomass and abundance were evenly distributed in the Yueqing Bay in November. Moreover, there was marked seasonal variation in the species diversity of zooplankton, which conformed to the abundance of zooplankton. Results of the dilution experiments indicated that there was grazing pressure of microzooplankton on phytoplankton in the Yueqing Bay throughout the year though the rate of microzooplankton grazing on phytoplankton varied seasonally. Phytoplanktons were growing at 0.26–2.07/d and grazed by microzooplankton at a rate of 0.15–0.48/d in different seasons. __________ Translated from Acta Ecologica Sinica, 2005, 25(8): 1853–1862 [译自: 生态学报, 2005, 25(8): 1853–1862]     

Seasonal dynamics and grazing rate of zooplankton in Yueqing Bay,China

The species composition,biomass,abundance,and species diversity of zooplankton were determined for samples collected from August 2002 to May 2003 from 14 stations in Yueqing Bay,China.Phytoplankton growth rate and microzooplankton grazing rate were obtained by using the dilution method developed by Landry and Hassett.The spatial and temporal variations of zooplankton and its relationship with environmental factors were also analyzed.The results showed that the zooplankton in the Yueqing Bay could be divided into four ecotypes,namely coastal low saline species,estuary brackish water species,offshore warm water species,and eurytopic species.A total of 75 species of zooplankton belonging to 56 genera and 17 groups of pelagic larva were identified in the Yueqing Bay.The coastal low saline species was the dominant ecotype in the study area,and the dominant species were Labidocera euchaeta,Acartia pacifica,Acrocalanus gibber,Pseudeuphausia sinica,and Sagitta bedoti among others.There was considerable seasonal variation in zooplankton biomass and abundance in the surveyed areas.The peak biomass appeared in August,descending in November and in May,and the lowest biomass appeared in February.Similarly,the highest abundance of zooplankton was observed in August,with the abundance descending in the following months:May,November,and February.There were similar horizontal distribution patterns for the biomass and the abundance of zooplankton.They both increased from the upper to the lower bay in February and May,but decreased from the upper to the lower bay in August.Biomass and abundance were evenly distributed in the Yueqing Bay in November.Moreover,there was marked seasonal variation in the species diversity of zooplankton,which conformed to the abundance of zooplankton.Results of the dilution experiments indicated that there was grazing pressure of microzooplankton on phytoplankton in the Yueqing Bay throughout the year though the rate of microzooplankton grazing on phytoplankton varied seasonally.Phytoplanktons were growing at 0.26-2.07/d and grazed by microzooplankton at a rate of 0.15--0.48/d in different seasons.     

Effect of inorganic nutrients on the acclimation period preceding mineralization of organic chemicals in lake water

The addition of phosphate, nitrate, or sulfate (each at 10 mM) decreased the acclimation period for the mineralization of low concentrations of p-nitrophenol (PNP) in lake water. Added phosphate shortened the acclimation period for biodegradation of 2 ng to 2 micrograms of PNP per ml in various lake water samples and of 2,4-dichlorophenoxyacetate at 100 ng/ml. Added P enhanced the rate of growth of PNP-mineralizing microorganisms in waters containing 200 ng or 2 micrograms of PNP per ml. We suggest that the effect of P on the acclimation period results from an increase in the growth rate of the initially small population of microorganisms able to mineralize the synthetic chemicals.     

Combined effects of zooplankton grazing and dispersal on the diversity and assembly mechanisms of bacterial metacommunities

Effects of dispersal and the presence of predators on diversity, assembly and functioning of bacterial communities are well studied in isolation. In reality, however, dispersal and trophic interactions act simultaneously and can therefore have combined effects, which are poorly investigated. We performed an experiment with aquatic metacommunities consisting of three environmentally different patches and manipulated dispersal rates among them as well as the presence or absence of the keystone species Daphnia magna. Daphnia magna reduced both local and regional diversity, whereas dispersal increased local diversity but decreased beta‐diversity having no net effect on regional diversity. Dispersal modified the assembly mechanisms of bacterial communities by increasing the degree of determinism. Additionally, the combination of the D. magna and dispersal increased the importance of deterministic processes, presumably because predator‐tolerant taxa were spread in the metacommunity via dispersal. Moreover, the presence of D. magna affected community composition, increased community respiration rates but did not affect bacterial production or abundance, whereas dispersal slightly increased bacterial production. In conclusion, our study suggests that predation by a keystone species such as D. magna and dispersal additively influence bacterial diversity, assembly processes and ecosystem functioning.     

A new sensitive tracer for the determination of zooplankton grazing activity

A new tracer compound is presented for determining zooplanktongrazing activity. The gut content in zooplankton is measuredas ß-dimethylsulphoniopropionate (DMSP), which canbe measured even in individual copepods. Species-specific DMSP/Chl-aratios allow applications in, for example, prey selectivitystudies.     

The effect of zooplankton grazing on estuarine blooms of the toxic dinoflagellate Gonyaulax tamarensis

A series of short-term ^{in situ} experiments was conducted intwo Cape Cod embayments to estimate mortality rates of the toxicdinoflagellate, Gonyaulax tamarensis, resulting from grazingby zooplankton. Rates of grazing by the whole zooplankton communityand by specific zooplankton populations were measured at variouspoints in the G. tamarensis bloom cycle. The planktonic larvaeof the spionid polychaete Polydora ligni and the tintinnid ciliateFavella sp. were important grazers in the systems studied. Gonyaulax-specificclearance rates effected by Polydora ranged from 0.02 to 0.5ml individual^–1 h^–1; for Favella the range was aboutan order to magnitude lower. Peak population densities wereclose to 900 and 400 individuals 1^–1 for P. ligni andFavella, respectively. Whether measured directly or predictedas the product of individual clearance rates and numerical abundance,rates of grazing were often higher than estimated algal divisionrates in years when blooms failed to develop. A simulation modelcorroborated the results of the field study, demonstrating thatgrazing can be a significant source of mortality during blooms,and can suppress bloom development when grazers are abundant.     

Relative growth rates and the grazing optimization hypothesis

Summary A mathematical analysis of the changes in plant relative growth rates necessary to increase aboveground production following grazing was conducted. The equation derived gives an isoline where production of a grazed and ungrazed plant will be the same. The equation has four variables (mean shoot relative growth rate, change in relative growth rate after grazing, grazing intensity, and recovery time) and may be analyzed graphically in a number of ways.Under certain conditions, small increases in shoot relative growth rate following grazing will lead to increased aboveground production. Under other conditions, very large increases in relative growth rate after grazing can occur without production being increased over that of ungrazed plants. Plants growing at nearly their maximum potential relative growth rate have little opportunity to respond positively to grazing and potentially can sustain less grazing than plants with growth rates far below maximum. Plants with high relative growth rates at the time of grazing require large increases in growth rate while slow growing plants require only small increases. High grazing intensities are least likely to increase production and high grazing frequencies require greater responses than infrequent grazing events.     

Effect of inorganic nutrients on the acclimation period preceding mineralization of organic chemicals in lake water.

The addition of phosphate, nitrate, or sulfate (each at 10 mM) decreased the acclimation period for the mineralization of low concentrations of p-nitrophenol (PNP) in lake water. Added phosphate shortened the acclimation period for biodegradation of 2 ng to 2 micrograms of PNP per ml in various lake water samples and of 2,4-dichlorophenoxyacetate at 100 ng/ml. Added P enhanced the rate of growth of PNP-mineralizing microorganisms in waters containing 200 ng or 2 micrograms of PNP per ml. We suggest that the effect of P on the acclimation period results from an increase in the growth rate of the initially small population of microorganisms able to mineralize the synthetic chemicals.