Description and use of an improved method for determining estuarine zooplankton grazing rates on phytoplankton

A method of rapidly determining zooplankton grazing rates on natural mixed phytoplankton populations using ¹⁴C is described. The method simplifies the design of grazing experiments as the grazing time can be kept short enough to prevent recycling of the isotope, and growth of the phytoplankton substrate. Very high specific activity, ¹⁴C-labelled phytoplankton concentrated either by centrifugation or sieving, may be used either as the sole grazing substrate, or as a tracer in natural mixed phytoplankton. Zooplankton, confined in glass jars at either ambient, or higher than ambient concentrations, are permitted to feed on the phytoplankton for periods of 30 min and 2 h, and are then separated by sieving. The zooplankton community grazing rate, or, if the samples are sorted into species, the individual species grazing rates, can be determined after scintillation counting of the zooplankton. The rate of appearance of ¹⁴C-labelled phytoplankton in the zooplankton is an estimate of the grazing rate, and the slope of the line joining the grazing rates at various phytoplankton concentrations gives an estimate of the grazing rate constant for the zooplankton population. The method provides a quick way of obtaining both zooplankton population, and individual species grazing rates on natural mixed phytoplankton. In two experiments, labelled phytoplankton was used as the sole grazing substrate in concentrations ranging between 0.4 and 5 times ambient levels. Grazing rate constants, for net-caught zooplankton concentrated to 46 times (Experiment 1) and 28 times (Experiment, 2) ambient estuarine levels were-0.14and-0.12 of the phytoplankton standing stock per day, respectively. There was a linear increase in the amount of phytoplankton grazed with an increase in phytoplankton concentration up to four times ambient phytoplankton levels. When tracer amounts of labelled phytoplankton were added to samples containing both phytoplankton and zooplankton at ambient concentrations the grazing rate constants were-0.28 and-0.42 of the phytoplankton standing stock per day. We conclude that zooplankton grazing was the major control factor of phytoplankton population size during October–November 1975 in South West Arm, Port Hacking, near Sydney, Australia.     

Role of herbivory in controlling phytoplankton abundance: annual pigment budget for a temperate marine fjord

An annual pigment budget was constructed for Dabob Bay, Washington (USA) by comparing the downward vertical loss of phytoplankton pigments (chlorophyll and pheopigments) to the production of chlorophyll within the euphotic zone. The vertical flux of pigments was measured with sediment traps deployed at intervals of 1 to 6 wk over a 2.5 yr period yielding 763 d of trap exposure (28 November 1978–16 June 1981). The production rate of chlorophyll was calculated from measurements of algal specific growth rates, chlorophyll (chl) crops, primary production (as carbon) and appropriate C: chl ratios. Sixty one to 77% of the annual chlorophyll production was accounted for by the vertical flux of pheopigments resulting from herbivorous zooplankton grazing (macrozooplankton). Algal sinking, represented by downward chlorophyll flux, accounted for only 5 to 6% of the annual chlorophyll production. The remaining fraction of chlorophyll production was estimated to be consumed by small herbivores (microzooplankton), whose fecal material contributes to the suspended pool of pheopigments found in the euphotic zone. The suspended pheopigments are continuously removed by photodegradation. In Dabob Bay, the major process controlling phytoplankton abundance is zooplankton grazing and it appears that the ultimate fate of most phytoplankton is to be consumed by herbivores.     

Variability and its effect on the 24h chlorophyll budget of a small marine basin

In an intensive study (lasting 25 h) of the production, export and grazing of phytoplankton in a small marine basin, it was found that 58% of the production (11% of the total standing stock) was lost by exchange with the sea and 34% was consumed by grazing of zooplankton. The measured production of phytoplankton could be balanced, to within a few percent, against grazing, export, and a small, measured, net change in the total standing stock of the basin. Large variations were observed in concentrations of chlorophyll and zooplankton at the mouth of the basin over the 25 h period. These variations were associated with changes in the height of the tide, but were about 2^1/2 h out of phase with it. Strong negative correlations were observed between chlorophyll and transport, such that only 35% of the chlorophyll exported was exchanged via the mean flow, while 65% was exchanged via the fluctuations. The correlation was even more striking with zooplankton, for which virtually all the export was associated with the fluctuations in the transport. Time series observations in the centre of the basin revealed considerable short-term variability in both chlorophyll and zooplankton, but the variations were smaller than those observed at the mouth of the basin, and the phase lag with the tide was longer. The variability studies enable suggestions to be made about more economical design of sampling programs, but illustrate the difficulty of providing verification data for any continuous model of primary production in such a basin.Bedford Institute Contribution No. 231.Canadian Contribution to IBP No. 97.     

Zooplankton biomass and indices of grazing and metabolism during a late winter bloom in subtropical waters

The development of the so-called late winter bloom in subtropical water was studied in an oceanic area north of the Canary Islands from January to May 2000. Zooplankton was sampled at short-term intervals (1–4 days) during the bloom (January–March), and biomass, indices of grazing (gut fluorescence) and metabolism (electron transfer system activity, ETS) were measured in four different size fractions (100–200, 200–500, 500–1000 and >1000 µm). During the bloom, ETS activity and gut fluorescence increased before the development of zooplankton biomass. At the end of February, the presence of an impressive cloud of dust formed in the Sahara desert was related to an increase in chlorophyll and small zooplankton a week later. The increments in biomass were the consequence of consumption by zooplankton as inferred from the indices of grazing and metabolism. Estimated grazing from gut fluorescence and gut evacuation rates during the period of study accounted for 55% of the assessed total ingestion from respiration and normal values of assimilation, showing the importance of the non-pigmented food in the diet of zooplankton in these waters. In contrast, the sharp decreases in zooplankton biomass observed during the bloom appeared during the dark period of the moon, the days in which the diel vertical migrants reach the shallower layers, in agreement with previous works in the area. Thus, the development of the late winter bloom in this region is suggested to be driven by the interplay between resource and consumer controls.Communicated by S.A. Poulet, Roscoff     

In situ grazing rates of deep-sea benthic boundary-layer zooplankton

In situ grazing rates of mixed assemblages of deep-sea benthic boundary-layer zooplankton were measured in July 1983 at a site in the North Atlantic Ocean at 2 175 m depth using a short-term radioisotope-uptake method. Zooplankton were collected with an opening-closing net system from the bottom 1 m of the ocean and incubated in situ with mixed tritiated amino acids in special cod-end chambers. Incubations were terminated at depth by the addition of MS-222. Radioisotope uptake beyond that of dead controls was shown by both the zooplankton and particulate fractions. Grazing rates in the deep-sea experiments were surprisingly high, being comparable to wintertime Narragansett Bay zooplankton grazing rates determined in a separate series of laboratory experiments. These laboratory experiments also documented nonparticle-associated uptake of dissolved amino acids by Narragansett Bay zooplankton, but the importance of this in the deep sea is unknown. The deep-sea benthic boundary layer may be a region of elevated rates and activity because of its higher particulate concentrations, and our experiments may also have measured maximum rather than average rates.     

Effects of food-level acclimation on digestive enzyme activities and feeding behavior of Calanus pacificus

A recent hypothesis in the zooplankton literature states that zooplankton acclimate to ambient food concentrations such that higher digestive enzyme activities and, consequently, higher maximum ingestion rates are achieved at higher food levels. To test this hypothesis, adult female Calanus pacificus, collected from the main basin of Puget Sound, Washington, USA, in August 1979 and May 1982, were conditioned for 2 wk at different concentrations of the diatom Thalassiosira weissflogii (=fluviatilis). Ingestion rates and the activity of the digestive enzymes laminarinase, maltase, and cellobiase were measured periodically during acclimation and in a block-designed feeding experiment at the end of acclimation. Consistent with the hypothesis, maximum ingestion rate and digestive enzyme activity were positively correlated. However, in contrast to the hypothesized mechanism, this result arose because both maximum ingestion rate and digestive enzyme activity were negatively correlated with food concentration during acclimation. The enhanced ingestion of copepods following long-term (12 to 14 d) acclimation to low food is similar to that previously described for short-term (e.g. 1 d) starvation. It might be energetically optimal for copepods experiencing a patchy food environment to maintain higher levels of digestive enzymes at low food concentrations in order to exploit high concentrations of food when encountered.     

Spring and autumn spatial distribution of zooplankton carbon requirement across the Mediterranean Sea

Zooplankton represents a key contributor to the ocean biological pump through its consumption of sinking and suspended carbon. A specific and highly sensitive method to evaluate zooplankton carbon requirement from the sinking flux is through the estimation of the activity of the electron transport system. The present study was carried out from samplings in 2006, and it was focused on the spatial 200–0?m zooplankton carbon demand across 24 sampling stations, along the Mediterranean Sea, from the island of Crete to the Strait of Gibraltar. Its potential day/night variability was evaluated. The zooplankton composition, abundance and biomass were investigated. The carbon demand per unit zooplankton biomass indicates geographical and diel differences among the sampling stations. A higher mean carbon demand was seen for the western Mediterranean with respect to the eastern Mediterranean, which can be justified through the observed ratio of gelatinous:crustacean taxa and the water temperatures recorded. Higher carbon demand was measured in samples collected during the dark hours. The relation to the presence and abundance of actively migrating euphausiids and copepods was discussed. A comparison with data from another of our study carried out in the same study area but in another seasonal period was done.     

Phytoplankton carbon dynamics during a winter-spring diatom bloom in an enclosed marine ecosystem: primary production,biomass and loss rates

Phytoplankton production, standing crop, and loss processes (respiration, sedimentation, grazing by zooplankton, and excretion) were measured on a daily basis during the growth, dormancy and decline of a winter-spring diatom bloom in a large-scale (13 m³) marine mesocosm in 1987. Carbonspecific rates of production and biomass change were highly correlated whereas production and loss rates were unrelated over the experimental period when the significant changes in algal biomass characteristic of phytoplankton blooms were occurring. The observed decline in diatom growth rates was caused by nutrient limitation. Daily phytoplankton production rates calculated from the phytoplankton continuity equation were in excellent agreement with rates independently determined using standard ¹⁴C techniques. A carbon budget for the winter bloom indicated that 82.4% of the net daytime primary production was accounted for by measured loss processes, 1.3% was present as standing crop at the end of the experiment, and 16.3% was unexplained. Losses via sedimentation (44.8%) and nighttime phytoplankton respiration (24.1%) predominated, while losses due to zooplankton grazing (10.7%) and nighttime phytoplankton excretion (2.8%) were of lesser importance. A model simulating daily phytoplankton biomass was developed to demonstrate the relative importance of the individual loss processes.     

Day-night differences in the emergence of demersal zooplankton from a sand substrate in a kelp forest

Day-night differences in abundance and biomass of demersal zooplankton in the water column were determined by trapping these animals as they emerged from the sand substrate in a kelp forest (Macrocystis pyrifera) ecosystem off Santa Catalina Island, California, USA. The day and night sampling periods of the 24 June 1979 new moon each lasted 12 h. Abundance and biomass of total demersal zooplankton were significantly higher in night samples. A mean of 2,425±1,168 demersal zooplankton m^-2 24 h^-1 migrated over a diel cycle; 97% of these animals were crustaceans. The mean biomass of demersal zooplankton was 94.2±27.6 mg ash-free dry wt m^-2 24 h^-1. No significant differences were found in either the abundance or biomass of demersal zooplanktion collected in low and high traps, suggesting that most animals collected 25 cm off the bottom can sustain swimming to at least 75 cm and that both traps give comparable estimates of the amount of demersal zooplankton available to planktivorous predators.     

Zooplankton abundance and grazing at Davies Reef,Great Barrier Reef,Australia

Zooplankton abundance and grazing on autotrophic and heterotrophic particulate matter were measured along a transect across Davis Reef (18°5S; 147°39E) and in the back-reef lagoon over tidal and diel cycles during austral winter (August 1984). Zooplankton entering the reef from the surrounding shelf waters decreased in abundance over the reef flat, presumably because of predation. Within the reef lagoon, maximum daytime densities of pelagic copepods occurred during high water, suggesting an external input. At night, water-column zooplankton biomass increased by a factor of 2 to 3 due to the emergence of demersal reef zooplankton. Zooplankton grazing rates on heterotrophic particulate matter (bacteria + detritus and Protozoa) compared to phytoplankton were higher on the reef flat than on the fore-reef or lagoon. Within the lagoon, zooplankton grazing rates on heterotrophic material were maximum during high water, coincident with maximum tidal concentrations of particulate organic carbon. The combined demersal and pelagic zooplankton community were often able to crop 30% of the daily primary production by >2µm phytoplankton. However, >50% of phytoplankton biomass was in cells <2µm, presumably unavailable to these zooplankton. Our particulate production and ingestion measurements, together with zooplankton carbon demand extrapolated from respiration estimates, suggest that the zooplankton community of Davies Reef derives much of its nutrition from detritus.Joint contribution from the University of Maryland, Center for Environmental and Estuarine Studies (No. 2015), and the Microbial Ecology on a Coral Reef Workshop (MECOR No. 19)     

A method to determine in situ zooplankton grazing rates on natural particle assemblages

In situ zooplankton grazing rates on natural particle assemblages were stimated by measuring zooplankton uptake of labelled autotrophic (with Na¹⁴CO₃) and heterotrophic (with [methyl-³H]-thymidine) particulate matter in 1-h incubations in clear, Plexiglas, Haney chambers. The in situ grazing rates are in the same range as those measured for zooplankton in the laboratory using standard particle counting techniques. A negative selection coefficient for ³H-labelled particles indicated a lower filtration efficiency or avoidance of these particles by zooplankton.     

Influence des conditions hivernales sur les productions phyto-et zooplanctoniques en Méditerranée Nord-Occidentale. V. Biomasse et production zooplanctonique — relations phyto-zooplancton

The cruise “Mediprod I” of the R.V. “Jean Charcot” covered an area of the Mediterranean Sea situated North of the 40th parallel and West of Corsica; during two 15-day legs, the first when Winter conditions ended, in March, the second in Spring conditions, in April, a 48-station network was surveyed as to primary and secondary production, as well as hydrological conditions. The first survey revealed a higher concentration of zooplankton in hydrologically stable areas, especially in the South-East, although zooplankton values were rather low throughout the whole area surveyed. The total zooplankton per unit surface was lower at the edges of the surveyed area, where phytoplankton was scarce. An increase in zoo-plankton biomass was observed between the two surveys, mostly in the central areas and near the surface. An important difference between both legs is in the proportion of organisms of different sizes collected by the Clarke-Bumpus sampler: the-200 μ:+200 μ organisms ratio, which is around 1 during the first leg, is much lower during the second leg. Two methods were used in estimating the biomass: the +200 μ fraction of a Clarke-Bumpus sample (Cl-B towed with a 50-μ net fitted on the sampler) was collected by sieving the sample through a 200-μ mesh nylon; standard vertical WP 2 hauls were performed (200 μ mesh). Both roughly show the same zooplankton (weight per unit surface) distribution pattern. However, higher estimations of the total biomass of the volume investigated were sometimes provided by the Clarke-Bumpus method during the first leg, probably due to the distribution of the animals according to their size-classes. A graph of chlorophyll versus zooplankton for surface waters suggests that zooplankton has a limiting effect on the development of phytoplankton in April only. Apparent growth rates of cooplankton are less than those for populations in the laboratory or enclosed environments. Values of the mean secondary production vary from 18 mg C.m^-2.day^-1 for the first leg, to 230 mg C.m^-2.day^-1 for the second leg. Estimations of net efficiency for energetic transfer between phyto- and zooplankton lie between 7 and 26%. As far as our hypotheses as regards physiological coefficients are valid, we can assume that the effect of grazing relative to primary production is greater in the border areas than in the central area, thus increasing the contrast between both areas with time. Phosphorus excretion rate by zooplankton seems to be less than that measured in the Atlantic Ocean. We suggest that in Spring zooplankton excretion is not the main phosphorus recycling process. Primary productivity measurements, apparent growth coefficients, and estimated grazing rate have been used to calculate the expected mean biomass per unit area during the second leg. A 7% loss of phyto- and zooplankton from the upper 100-m layer must be assumed to explain the observed biomass variation. Vertical mixing and sinking of surface water on an isopycnal slope are responsible for such loss, which can also affect, to an undetermined degree, the phosphorus stock introduced into the surface layer by the Mediterranean deep-water formation mechanism. We suggest that nearly half the total loss of phytoplankton from March to April is attributable to animal grazing.     

The relationship between lunar periodicity and activity of fish-parasitic gnathiid isopods in the Caribbean

Gnathiid isopod larvae are members of the marine demersal zooplankton community and are common ectoparasites of coral reef fishes, emerging from the substratum, mostly at night and crepuscular periods to feed on fish blood. Given that the activity of many marine organisms is often linked to changes in the phase of the moon, we examined the relationship between lunar phase and activity in gnathiid isopods on Caribbean reefs. We employed two sampling methods to quantify gnathiid activity: (1) Emergence traps set on reefs over a 24 h period; and (2) live fish placed in cages on reefs and retrieved during night and dawn peaks in gnathiid activity. These were compared during discrete phases as well as a continuous metric, the lunar “angle”, and an estimate of ambient light availability (luminance). Lunar phase and angle varied in their statistical effect on gnathiid activity patterns by sampling method and/or year. Luminance had a significant but inconsistent effect on measures of gnathiid activity. Our results suggest that changes in the lunar cycle are not a strong predictor of gnathiid activity at our shallow reef study sites.     

Effects of mercury on the behaviour and oxygen consumption of Monodonta articulata

The trochid snail Monodonta articulata Lamarck was exposed to mercuric sulphate at concentrations of 0.2, 0.5, 0.8 and 1 ppm Hg⁺⁺. At 24 h, retraction into the shell was observed in 0.8 and 1 ppm Hg⁺⁺; this retraction increased in these concentrations at 36 h. Retracted snails died if retained in the solutions, but generally recovered within 24 to 48 h if transferred to uncontaminated sea water. Immersionemersion behaviour and interface activity were studied over 24 h by means of an aktograph; snails in normal sea water spent more time below than above the water surface, and exhibited frequent periods of activity. Exposure to mercuric sulphate at concentration of 0.25, 0.5, 0.8 and 1 ppm Hg⁺⁺ progressively reduced both the length and frequency of activity periods. From 0.5 ppm Hg⁺⁺ upwards, emersion periods increased, and immersion periods decreased. Oxygen consumption of snails was measured in sea water and in mercuric sulphate at concentrations of 0.2, 0.5, 0.8 and 1 ppm Hg⁺⁺. Oxygen consumption decreased significantly with each progressive rise in mercury concentration. It is considered that mercury affects M. articulata by interfering with respiration, initially reducing interface activity, then forcing the snail for longer and longer periods out of the water. Retraction occurs when activity is no longer possible. It is concluded that respiratory and behavioural alterations of this nature would afford a good indicator of the presence of sub-lethal concentrations of pollutants.     

Water-soluble vitamins in natural plankton (copepods) during two consecutive spring blooms compared to vitamins in Artemia franciscana nauplii and metanauplii

Weight-specific contents of asorbic acid, thiamin, riboflavin, niacin, pantothenic acid, vitamin B₆, biotin, folate and vitamin B₁₂ were analysed in natural zooplankton collected from a sea water lagoon in a halibut fry production plant. The samples of natural zooplankton were collected from April/May to the end of June during two consecutive spring blooms. For comparison, the same vitamins were measured in nauplii of Artemia franciscana, just after hatching and after enrichment for 24 and 48 h with a commercial diet. The changes in vitamin content in zooplankton, which consisted mainly of Temora longicornis, during the two spring blooms were <2.5-fold for each specific vitamin. The large number of high quality halibut fry produced using the plankton as a dietary source suggests that the plankton contained sufficient amounts of all water-soluble vitamins. With the exception of thiamin and ascorbic acid, Artemia– both newly hatched and enriched for 24 and 48 h – contained higher levels of water-soluble vitamins than natural zooplankton. Enrichment feeding of Artemia for 48 h stabilised or increased the level of the water-soluble vitamins. Received: 8 March 1999 / Accepted: 21 February 2000     

Pigment composition and size distribution of phytoplankton in a confined Mediterranean salt marsh ecosystem

Pigment composition and size distribution of phytoplankton were analysed in a group of Mediterranean salt marshes, where hydrology is dominated by sudden inputs during sea storms, followed by long periods of confinement. These marshes are characterized by a low inorganic–organic nutrient ratio, and inorganic nitrogen is especially scarce due to denitrification. Nutrients were the main factor affecting phytoplankton biomass, while zooplankton grazing did not control either phytoplankton community composition, or their size distribution. The relative abundance of the different phytoplankton groups was analysed by correspondence analysis using the pigment composition measured by high-performance liquid chromatography (HPLC) and analysed with the CHEMTAX programme. In this analysis, phytoplankton pigment composition was correlated with two nutrient gradients. The first gradient was the ratio of nitrate–total nitrogen (TN), since the different phytoplankton groups were distributed according to their eco–physiological differences in nitrogen uptake. The second gradient was correlated with total nutrient loading. Biomass size distributions frequently showed a lack of intermediate sized nanophytoplankton (2.5–4 μm in diameter), and the importance of this lack of intermediate sizes correlated with dinoflagellate biomass. These results suggested that in confined environments, where nutrients are mainly in an organic form, dinoflagellates take advantage of their mixotrophy, by competing and grazing on smaller phytoplankters simultaneously.     

Short-term changes in feeding and digestion by the copepodCalanus pacificus

The planktonic marine copepodCalanus pacificus exhibits an enhanced feeding rate, or hunger response, when exposed to food following short periods of starvation. In a scries of laboratory experiments with copepods collected from the main basin of Puget Sound, Washington, during 1982 and 1984, we measured maximum ingestion rate, assimilation efficiency, and digestive enzyme activity to determine the time scales over which the feeding behavior ofC. pacificus responds to increases in food. These laboratory results were then compared to field studies of diel fluctuations in digestive enzymes and gut fluorescence ofC. pacificus in Dabod Bay, a fjord of Puget Sound, during September, 1980, and the closely relatedC. marshallae off the Washington coast, in August, 1981. Laboratory experiments demonstrated that the hunger response ofC. pacificus lasts approximately 6 h before ingestion rate returns to a steady state level of about one-half maximum. On the order of 12h of starvation were required to induce the maximum ingestion rate of the hunger response. Digestive enzyme activities did not change over these time scales. Assimilation efficiency peaked within a few hours of the onset of feeding, with low initial rates possibly related to the period of starvation prior to feeding. These results were consistent with diel patterns observed in the field. The hunger response ofC. pacificus appears to be controlled by processes within the gut, and our results are discussed in relation to recent studies of the digestive processes of calanoid copepods.Contribution No. 1772 from the School of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Temporal variability of zooplankton biomass (ATP content and dry weight) in the St. Lawrence Estuary: Advective phenomena during neap tide

In order to assess the temporal variability of living zooplankton in a tidal estuary, the ATP content and dry weight of mixed zooplankton populations (mainly copepods) were measured during a period of 175 h at an anchor station in the Upper St. Lawrence Estuary. Vertical tows were made every 30 min. Hourly vertical profiles of the current speed and direction, temperature and salinity were also obtained during the experiment. A strong tidal influence was found in all series. Maxima and minima of the ATP content (living biomass) and the dry weight (total biomass) were correlated with low and high water slacks. The serial autocorrelation and cross-correlation showed, in both series a 12 to 13-h cycle, and the ATP:dry weight ratio showed a significant 24-h cycle. The cross-correlation with the Kendall τ was used to detect the relationship between biological components and physical indices (stratification and Ri). It is suggested that the proportion of living zooplankton biomass in the Upper St. Lawrence Estuary is most likely the result of a combination of diurnal migration and longitudinal advection.     

Nitrogen excretion by some demersal macrozooplankton in Heron and One Tree Reefs,Great Barrier Reef,Australia

Nitrogen excretion rates of demersal macrozooplankton were measured together with nitrogen concentrations in the water column and sediments in lagoons of Heron Reef and One Tree Reef, Great Barrier Reef, Australia, during August and November 1991. Excretion rates increased with body weight, and weight-specific excretion rates of the demersal macrozooplankton were comparable to those of pelagic zooplankton and meiofauna in the Great Barrier Reef. Values of demersal macrozooplankton abundance from previous studies and excretion rates from this study were combined to estimate fluxes of ammonium from demersal macrozooplankton in coral reef lagoons. The estimated fluxes in the water column and sediments were 12 M NH₄ m^-2 d^-1 and 34 M NH₄ m^-2d^-1, respectively. These fluxes were compared with reported fluxes of ammonium in coral reef lagoons in the Great Barrier Reef, Australia. The estimated flux from the demersal macrozooplankton in the water column was 29 and 9% of those reported for microheterotroph regeneration and phytoplankton utilization, respectively. It was 10% of the reported advective flux during periods of low advection and 13% of the maximum efflux from sediments computed from diffusion models. The estimated flux from the demersal macrozooplankton in the sediments exceeded those reported for meiofauna, and was 5 to 32% and 2 to 13% of those reported for ammonification and utilization in sediments, respectively. The potential importance of demersal macrozooplankton in mediating sediment-water column exchanges in the absence of diffusive effluxes and when they swarm is discussed.     

Effects of zooplankton diel vertical migration on a phytoplankton community: A scenario analysis of the underlying mechanisms

A mechanistic model was applied to study the influence of diurnal vertical migration (DVM) of planktonic crustaceans on the succession and composition of the phytoplankton community. While zooplankton was restricted to only one functional group, the phytoplankton community was divided into two functional groups which are distinguished by their maximum growth rates and vulnerability to zooplankton grazing. DVM causes a pulsed grazing regime and may also entail a corresponding reduction of the cumulative daily rates of ingestion and losses of zooplankton. To study the relative importance of these two mechanisms of DVM to phytoplankton we performed a scenario analysis consisting of 5 different scenarios. The results show that DVM has a strong influence on the phytoplankton community. Well edible algae benefit during the first 3–4 weeks of summer stratification by reduced daily grazing. The typical shift from small, well edible algae to larger, poorly or non-edible phytoplankton is distinctly delayed. Under the assumption of unchanged daily grazing, however, a pulsed grazing regime has nearly no influence on the resulting phytoplankton composition. As similar effects are also found for completely non-edible phytoplankton, indirect effects via phosphorus availability must be assumed. Thus, the scenario analysis reveals that the observed effects of DVM on phytoplankton can be explained by a combination of two mechanisms: (1) reduction of the daily zooplankton grazing, and (2) changed assimilation and remineralisation of phosphorus. Surprisingly and in contradiction to earlier reports there is almost no DVM effect on phytoplankton due to the sole action of a pulsed grazing regime.