Summer dynamics of the deep chlorophyll maximum in Lake Tahoe

Vertical profiles of chlorophyll and phytoplankton biomass weremeasured in Lake Tahoe from July 1976 through April 1977. Adeep chlorophyll maximum (DCM) persisted during summer and earlyautumn (July—October) near 100 m, well below the mixedlayer and at the upper surface of the nitracline. The DCM coincidedwith the phytoplankton biomass maximum as determined from cellcounts. In addition, the composition of the phytoplankton assemblagewas highly differentiated with respect to depth. Cyclotellastelligera was the predominant species in the mixed layer whilethe major species in the DCM layer included C. ocellata andseveral green ultraplanktonic species. In situ cell growth playsa substantial role in maintaining the DCM, but sinking of cellsfrom shallower depths and zooplankton grazing above the DCMmay contribute to the maintenance of the DCM. Calculations supportthe interpretation that the summer DCM persists at the boundarybetween an upper, nutrient-limited phytoplankton assemblageand a deeper, light-limited assemblage.     

Pigment ingestion and egg production rates of the calanoid copepod Temora longicornisr. implications for gut pigment loss and omnivorous feeding

We examined the relationship between egg production rate (E)and pigment ingestion rate (I, from gut content corrected for33% loss) for adult female Temora longicornis in Long IslandSound on 47 occasions. Linear regression of E on I [both variablesexpressed in mass of nitrogen (N) female^–1 day^–1]was: E_N = 0.0016 + 0.770 x I_N. The slope, 0.77, is the apparentgross efficiency of egg production, equivalent to the grossgrowth efficiency (GGE) assuming that females partition allnitrogen for growth into egg production. Published work suggeststhat a GGE of 0.37 would be expected for herbivorous copepods.The discrepancy between the expected value of 0.37 and observedvalue of 0.77 could result from unquantified losses of gut pigmentor because T.longicomis ingested a significant amount of nitrogenby feeding as a carnivore. We suggest that if T.longicomis femalesderive all of their nitrogen for growth by feeding on phytoplankton,and if no correction for pigment loss is employed, then thegut pigment method underestimates pigment ingestion by no morethan a factor of two.     

Influence of phytoplankton composition and stratification degree on gut pigment content of Ceriodaphnia sp. at dawn

Very short-term feeding activity of the cladoceran Ceriodaphniasp. was investigated in situ in a eutrophic reservoir in thesouth of Spain, using fluorimetric analysis of the gut pigmentcontent in periods when the water column was relatively mixedor strongly stratified. The results obtained in the mixed watercolumn showed a clear increase in gut pigment content at dawn,a period sampled with high frequency. The accumulation of thecladoceran at the depth of maximum concentration of phytoplankton,and the high gut pigment concentration in cladocerans at thatdepth just after dawn, suggested active feeding of Ceriodaphniaon phytoplankton at that time. During stratification, the abundanceof Ceriodaphnia was higher, but the gut pigment contents werevery low and they did not reflect any clear feeding patterns,with either time or depth. Changes in phytoplankton concentrationand composition between the relatively mixed and the stratifiedwater column suggest a shift in feeding activity from herbivorousto.     

Diel variations in gut pigment content, diel vertical migration and estimates of grazing impact for copepods in the southern Benguela upwelling region in October 1987

The gut fluorescence technique was used to estimate ingestionand filtration rates of the adult female copepods Paracalanusparvus, Cenlropages brachiatus and Calanus austrails, and copepoditestages 3, 4 and 5 of C.australis in the southern Benguela upwellingregion. During the study period chlorophyll concentrations withinthe upper 20 m of the water column were high, 5 µg I^–1in mid-shelf waters and 15–30 µg I^–1 in innershelf waters. Copepod gut pigment content was low and constantduring the day then increased sharply during the first 2 h aftersunset. Gut pigment content was 2–6 times higher duringthe night compared with daytime values. Small non-migratingcopepods (Paracalanus parvus) showed the smallest diel differencein gut pigment content and large migrating copepods (Centropagesbrachiatus and Calanus australis) the largest difference. Eggproduction rates were 20 and 50% of maximum at the mid-shelfand inner shelf stations respectively, suggesting food-limitation.Comparison of ingestion rates calculated from egg productiondata with ingestion rates calculated from gut pigment data suggestedthat the copepods were feeding omnivorously at the inner shelfstations but herbivorously at the mid-shelf stations. Assumingthat all of the phytoplankton was available as food, the nearshorecopepod assemblage grazed {small tilde}1% of the standing cropeach day, and the mid-shelf assemblage grazed 5% day^–1.Because of errors and uncertainties associated with the gutfluorescence technique, the feeding impact could be underestimatedby 2–4-fold. We discuss several approaches which couldlead to more precise estimates of feeding rates. ³Present address: Marine Sciences, SUNY, Stony Brook, NY, 11794-5000,USA     

Interannual variability in the distribution of the phytoplankton standing stock across the seasonal sea-ice zone west of the Antarctic Peninsula

The spatial distribution of phytoplankton cell abundance, carbon(C) biomass and chlorophyll a (Chl a) concentration was analysedduring three summers (1996, 1997 and 1999) in a seasonal sea-icearea, west of the Antarctic Peninsula. The objective of thestudy was to assess interannual variability in phytoplanktonspatial distribution and the mechanisms that regulate phytoplanktonaccumulation in the water column. Phytoplankton C biomass andChl a distributions were consistent from year to year, exhibitinga negative on/offshore gradient. The variations in C concentrationhad a close and non-linear relationship with the upper mixedlayer depth, suggesting that the vertical mixing of the watercolumn is the main factor regulating phytoplankton stock. Themagnitude of C gradients was 5-fold higher during 1996 thanduring 1997 and 1999. This was ascribed to interannual variationsin the concentration of diatom blooms in the region influencedby sea-ice melting. Vertical distribution of the phytoplankton,as estimated from Chl a profiles, also varied along an on/offshoregradient: Chl a was distributed homogeneously in the upper mixedlayer in coastal and mid-shelf stations and concentrated inthe deep layer (40–100 m) occupied by the winter waters(WW, remnants of the Antarctic surface waters during summer)in more offshore stations. The region with a deep Chl a maximumlayer (DCM layer) was dominated by a phytoplankton assemblagecharacterized by a relatively high concentration of diatoms.The extent of this region varied from year to year: it was restrictedto pelagic waters during 1996, extended to the shelf slope during1997 and occupied a major portion of the area during 1999. Itis hypothesized that iron depletion in near surface waters dueto phytoplankton consumption, and a higher concentration inWW, regulated this vertical phytoplankton distribution pattern.Furthermore, we postulate that year-to-year variations in thespatial distribution of the DCM layer were related to interannualvariations in the timing of the sea-ice retreat. The similaritybetween our results and those reported in literature for otherareas of the Southern Ocean allows us to suggest that the mechanismsproposed here as regulating phytoplankton stock in our areamay be applicable elsewhere.     

Fish-induced changes in zooplankton grazing on phytoplankton and bacterioplankton: a long-term study in shallow hypertrophic Lake Sobygaard

The impact of fish-mediated changes on the structure and grazingof zooplankton on phytoplankton and bacterioplankton was studiedin Lake Søbygaard during the period 1984–92 bymeans of in vitro grazing experiments (¹⁴C-labelled phytoplankton,³H-labelled bacterioplankton) and model predictions. Measuredzooplankton clearance rates ranged from 0–25 ml l^–1h^–1 on phytoplankton to 0–33 ml l^–1 h^–1on bacterioplankton.The highest rates were found during thesummer when Daphnia spp. were dominant. As the phytoplanktonbiomass was substantially greater than that of bacterioplanktonthroughout the study period, ingestion of phytoplankton was26-fold greater than that of bacterioplankton. Multiple regressionanalysis of the experimental data revealed that Daphnia spp.,Bosmina longirostris and Cyclops vicinus, which were the dominantzooplankton, all contributed significantly to the variationin ingestion of phytoplankton, while only Daphnia spp. contributedsignificantly to that of bacterioplankton. Using estimated meanvalues for clearance and ingestion rates for different zooplankters,we calculated zooplankton grazing on phytoplankton and bacterioplanktonon the basis of monitoring data of lake plankton obtained duringa 9 year study period. Summer mean grazing ranged from 2 to4% of phytoplankton production and 2% of bacterioplankton productionto maxima of 53 and 88%, respectively. The grazing percentagedecreased with increasing density of planktivorous fish caughtin August each year using gill nets and shore-line electrofishing.The changes along a gradient of planktivorous fish abundanceseemed highest for bacterioplankton. Accordingly, the percentagecontribution of bacterioplankton to the total ingestion of thetwo carbon sources decreased from a summer mean value of 8%in Daphnia-dominated communities at lower fish density to 0.7–1.1%at high fish density, when cyclopoid copepods or Bosmina androtifers dominated. Likewise, the percentage of phytoplanktonproduction channelled through the bacteria varied, it beinghighest (5–8%) at high fish densities. It is argued thatthe negative impact of zooplankton grazing on bacterioplanktonin shallow lakes is highest at intermediate phosphorus levels,under which conditions Daphnia dominate the zooplankton community.     

Diel periodicity of Subantarctic copepods: relationships between vertical migration, gut fullness and gut evacuation rate

A zooplankton community in the Polar Frontal Zone north of SouthGeorgia was sampled for 5 days in February 1994. Feeding ofvarious copepodite stages of six copepod species was assessedby a series of gut fluorescence/gut evacuation experiments.Feeding periodicity was compared to vertical distribution andmigration patterns, as revealed by Longhurst Hardy PlanktonRecorder (LHPR) and ring net catches. Despite chlorophyll alevels reaching only 0.8 mg m^–3 and daily carbon rationsbased on phytoplankton intake being low, feeding of all thecopepods was restricted mainly to the 8 h night-time perid.During the daytime, the epipelagic community was verticallydispersed within the top 100 m. At night, upward migration bymost species led to a convergence of almost all zooplanktersin the upper half of the surface mixed layer. However, large-amplitudevertical migrations which crossed the thermocline were performedonly by Metridia lucens and Pleuromamma robusta. Although feedingby both migrants and non-migrants was mainly at night, therewas no diel signal in gut evacuation rate. The gut evacuationrates of the nine copepod species and stages differed significantly(5-fold) and were negatively related to the extent of theirdiel vertical migration. The long gut-passage times of the migratoryspecies, M.lucens and P.robusta, would have allowed them timeto defaecate some of the food eaten near the surface at depth,contributing to an active carbon transfer out of the mixed layer.However, their scarcity at this site meant that their grazingcomprised only -<1.4% of total copepod ingestion, so theircombined gut flux is likely to have been negligible (-0.4 mgcarbon day^–1).     

The effects of food deprivation, prey density and volume on clearance rates and ingestion rates of Diacyclops thomasi

The ingestion rates of the copepod, Diacyclops thomasi, on thesoft-bodied rotifer, Synchaeta pectinata, increased 10-fold(0.07–0.77 Synchaeta h^–1) over the range 50–250prey l^–1. The saturating functional response curve appearedsigmoid but was statistically indistinguishable from a parabola.The response curve was more linear and 10 times lower over thesame range of density when Diacyclops was offered Kerarellacochleans, a species having a stiffened lorica. Diacyclops maximizedits ingestion rate on Synchaeta as a function of the availablegut space. Predation effort, measured as clearance rates, waslinked tentatively to changes in swimming speed of Diacyclopsand was a function of hunger level. Diacyclops, which were starvedfor varying periods of time, increased their ingestion rateson Synchaeta up to a maximum (-3.0 h^–1) after 7–10h of food deprivation. The gut passage time of Diacyclops wasestimated to be 7–8 h. Therefore, ingestion rates (andclearance rates) appeared to be strongly correlated to the volumeof food in the gut.     

Relationship between functional response and gut transit time in the calanoid copepod Acartia clausi: role of food quantity and quality

The relationship between ingestion rate and gut transit timeof the calanoid copepod Acartia clausi was examined in laboratoryexperiments with five different diets: (i) living cells of thediatom Thalassiosira weissflogii, (ii) detrital cells of thesame diatom, (iii) 50:50 mix of the two previous diets on aprotein basis, (iv) dinoflagellate cells of Prorocentrum micansand (v) Prorocentrum minimum. Ingestion followed a Holling type2 response for diets 1 and 4 and a linear one for diets 2, 3and 5. Gut transit time varied with food abundance only whenthe copepods were fed with the living diatom. The gut evacuationrate increased with the concentration of T. weissflogii withvalues of 0.010, 0.020, 0.032, 0.042 min^–1, correspondingto gut transit time of 97, 50, 31 and 24 min, measured at 50,110, 130 and 275 µg protein L^–1, respectively. Copepodsfed with dinoflagellates, mixed and pure detrital diets exhibitedlonger and similar gut transit times ranging from 85 to 166min, depending on diet. The coupling between ingestion rateand gut transit time measurements is discussed in the contextof copepod feeding strategies.     

A model for Acartia tonsa: effect of turbulence and consequences for the related physiological processes

A mathematical model of the energy budget of the coped Acartiatonsa is proposed. It takes into account the processes of ingestion(encounter, capture, ingestion sensu stricto), digestion (assimilation,gut transit and egestion), catabolic expenses and biomass production.In order to represent the potential effects of small-scale turbulenceon the whole physiological processes of a copepod, some processsubmodels already published are combined. A major assumptionof the model is a satiety effect resulting from midgut filling,which leads to a decrease in the feeding activity. The modelpermits the simulation of the short-term dynamics of ingestionunder different food and turbulence conditions, as well as anintegrated physiological balance over 24 h. The model is validatedthrough comparison with data at both scales Simulations showthat turbulence increases ingestion rates and gut contents,and causes a decrease in gut passage time and assimilation efficiency.As a consequence, the dependent physiological processes areaffected differently by turbulence, which preferentially increasesegestion and egg-production rates Simulated daily ingestionrates of A.tonsa, for suspension feeding on. Thalassiosira weissflogiiand for ambush predation on Strombidium sulcatum, are in goodagreement wth the available experimental observations. The concurrentdirect effect of turbulence on the copepod's metabolism, dueto increased escape reactions, is also simulated. Results ofthe model show that a switch from suspension feeding on diatomsin calm condition to ambush predation on ciliates in turbulentconditions, might allow A.tonsa to maintain its gross growthefficiency at the same level. It is suggested that a dynamicrepresentation of processes occurring over a time scale of afew seconds is necessary to obtain, once integrated over 24h, the correct simulation of the effect of microscale turbulenceon ingestion and the related physiological processes.     

The use of gut fluorescence to estimate grazing by oceanic salps

The fluorometric measurement of chlorophyll and its degradationproducts in the guts of salps has been used as a means of calculatingin situ grazing rates. Salps are particularly suitable for thismethod because they are non-selective filter-feeders large enoughto be individually collected by divers. Analysis of the pigmentin the gut has shown that chlorophyll degradation products canbe used as tracers to estimate in situ ingestion. Filteringrates are calcuLated from the amount of pigment, the gut clearancetime and the concentration of pigment in the water. For Pegeaconfoederata and Salpa maxima, gut-pigment rates are comparedwith rates measured in laboratory particle-clearance experiments.Salpa cylindrica has proved impossible to maintain in the laboratoryand the gut-pigment results are the first feeding data for thisspecies. Filtering rates for all species examined are high,on the order of liters per hour. Variability in the amount ofpigment found in S. cylindrica suggests this species, unlikethe others, may feed sporadically. The gut-pigment method permitsmeasurements of filtration and ingestion by herbivores on analmost instantaneous basis and can be a useful tool for assessinggrazing impact in field situations.     

A laboratory study of the effect of non-phytoplankton diets on the reproduction of Calanus helgolandicus

The effect of five non-phytoplankton and one phytoplankton dietson reproduction is described for the copepod Calanus helgolandicusand compared to field fecundity estimates. A fecundity increasefollowed ingestion of larvae of the sea urchin Sphaerechinusgranularis (LSU) and the oyster Crassostrea gigas (OYS) by copepodfemales, whereas a decrease followed ingestion of eggs of thecopepods Acartia and Temora spp. (COP), as a function of proteinconcentration in diets. With larvae of the sea urchin Paracentrotuslividus (SSU) and the dinoflagellate Prorocentrum minimum (PM;standard phytoplankton diet), fecundity was identical. Althoughprotein concentrations differed greatly from one food treatmentto the other, mean fecundity values were rather low. For unknownreasons, and despite their very high protein concentrationsin comparison to in situ, none of these diets could improvefemale fecundity or reach maximum egg production rates reportedin C.helgolandicus. Hatching rates were >80%, stable andsimilar with both PM and COP diets. With SSU, LSU and OYS diets,egg viability was lower than in situ and with PM, showing astrong and mild decrease with time with LSU and SSU, respectively.Ingestion of zygotes of the brown alga Fucus spiralis (ZYG)led to the death of females. These results suggest that denseblooms of zygotes of macroalgae and invertebrate embryos andlarvae, occurring in the continental shelf as part of copepoddiets, could have different effects on the demographic responseof C.helgolandicus.     

Natural ingestion rates and grazing impact of the marine cladoceran Penilia avirostris Dana in Tolo Harbour, Hong Kong

The gut fluorescence method was used to study algal grazingby Penilia avirostris in a semiestuarine embayment. Grazingimpact estimated from ingestion rate and animal abundance revealedthat only {small tilde}1% of the algal biomass was grazed perday by the cladocerans.     

Experimental evaluation of herbivory in the ctenophore Mnemiopsis leidyi relevant to ctenophore-zooplankton-phytoplankton interactions in Narragansett Bay, Rhode Island, USA

Herbivory of Mnemiopsis leidyi and its interactions with phytoplanktonand non-gelatinous zooplankton were examined in small-scalemicrocosm experiments. Clearance rates for M. leidyi incubatedwith phytoplankton were generally negative, but ranged up to4.5 1 ctenophore^–1 day^–1 when the large (80 µmø) diatom Ditylum brightwelli was offered as a food source.These highest ingestion rates would provide Mnemiopsis withonly 21 % of its daily carbon requirements for respiration.Mean shrinkage of M. leidyi was 8.2–51% when incubatedwith phytoplankton. Although M. leidyi neither fed activelyon phytoplankton, nor satisfied its nutritional needs on sucha diet, the chain-forming diatom Skeletonema costatum becameentangled in mucus strands and balls produced by M. leidyi inthe absence of zooplankton. Attachment onto mucus occurred atphytoplankton concentrations commonly observed in NarragansettBay and may be important in the formation of "marine snow" duringsummer M. leidyi pulses; phytoplankton sinking rate and the"package size" available to herbivores would also be affected.The experiments support our previous hypothesis based on fieldobservations in Narragansett Bay that M. leidyi indirectly regulatesphytoplankton abundance there during the summer bloom as a consequenceof predation on zooplankton. The extent to which M. leidyi influencedphytoplankton dynamics in the microcosms was dependent on therelative abundance and physiological state of the three trophiclevels. A food web diagram for M. leidyi is presented.     

Estimation of phosphorus metabolic rates of Daphnia galeata using a three-compartment model

A compartment model to estimate the different phosphorus metabolicrates in Daphnia galeata is presented. The model has three compartments:gut, metabolic pool and structural pool. Existing two-compartmentmodels used for carbon and phosphorus turnover in Daphnia donot allow estimation of ingestion and egestion rates. We extendedexisting two-compartment models with one more compartment, thegut, which allowed us to estimate both the ingestion and theegestion rates. Parameters of the model are estimated from asingle experiment of feeding unlabelled Daphnia with ³²P-labelledScenedesmus obliquus. Separate experiments with juvenile andadult daphnids were carried out in order to compare their metabolisms.This model permits a reliable estimation of the different metabolicrates of Daphrna in a single experiment and discriminates clearlybetween animals of different sizes.     

Cladoceran and rotifer grazing on bacteria and phytoplankton in two shallow eutrophic lakes: in situ measurement with fluorescent microspheres

Metazooplankton grazing on bacteria and on the phytoplanktonof various sizes was estimated in shallow eutrophic lakes Kaiavereand Võrtsjärv (Estonia) by in situ feeding experimentswith fluorescent microspheres (diameters 0.5 µm for bacteriaand 3, 6 and 24 µm for phytoplankton). Zooplankton communitycomposition, abundance and food density were important factorsdetermining grazing rates in these lakes. Cladocerans and rotifersfiltering rates (FR) and ingestion rates (IR) on bacteria andphytoplankton were several times higher in Lake Kaiavere wherebacterivorous rotifers and Daphnia contributed more to zooplanktonassemblage. While cladocerans were generally the main phytoplanktonconsumers, both lakes differed with respect to the groups ofbacterivores. Based on consumption of fluorescent microspheres,the metazooplankton grazing rates were relatively low and hadlow impact on production and standing stock of bacteria andingestible phytoplankton (<30 µm). On average, 0.5and 0.1% of standing stock of bacteria and 2.6 and 1.0% of standingstock of ingestible phytoplankton was grazed daily by metazooplanktonin lakes Kaiavere and Võrtsjärv, respectively. Thatcorresponded to daily grazing of 4.1% of the bacterial productionand 0.43% of the total primary production (PP) by metazooplanktonin Lake Kaiavere compared with 4.3 and 0.06% in Lake Võrtsjärv,respectively. The results suggest that the majority of consumptionof the bacterial and phytoplankton PP is most likely channelledthrough the microbial loop.     

Chlorophyll a conversion and gut passage time for the pelagic tunicate Oikopleura vanhoeffeni (Appendicularia)

We determined the chlorophyll (Chl) a conversion efficiency,defined as the conversion of Chi a into pheopigments and non-fluorescentproducts, for the cold-water appendicularian Oikopleura vanhoeffeniusing ⁶⁸Ge as conservative tracer. In both laboratory experimentsand in freshly collected animals, there was enough undegradedChi a present in the gut to provide an index of feeding activityfor individual animals. In laboratory experiments with diatoms,Chi a conversion efficiency was predictable with an averageof 79%. The mean gut passage time (0.8 h) determined using cornstarch and diatoms as markers was not influenced by food concentrationor animal size over a trunk length range of 1.8–5 mm.When experimentally determined Chi a conversion efficiency andgut passage times were applied to O.vanhoeffeni from Logy Bay(Newfoundland), the estimated clearance rates were in closeagreement with previously published values using several differenttechniques. We therefore suggest that the modified gut pigmenttechnique is a useful tool to assess in situ ingestion ratesof O.vanhoeffeni.     

Feeding efficiency of a marine copepod Acartia erythraea on eight different algal diets

Acartia erythraea is a dominant zooplankton copepod in the South China coastal waters during summer. This paper examined its feeding behavior (food gut passage, clearance rate and ingestion rate) on eight phytoplankton diets. The food gut passage time and ingestion rate were negatively related to the size and concentration of food supply, whereas the clearance rate was positively related to the food concentration. The ingestion rate decreased with the food concentration when it reached a threshold level. Generally, the clearance rate of copepods decreased with increasing cell density, but was very low at both low and high algal densities when the food were small in sizes. The optimum food size was about 10 μm for the copepods, and the dinoflagellate, Prorocentrum minimum, was considered as a good food choice for A. erythraea.     

Size-fractionated mesozooplankton biomass, metabolism and grazing along a 50{degrees}N-30{degrees}S transect of the Atlantic Ocean

Size-fractionated mesozooplankton grazing and metabolism wereinvestigated along the wide latitudinal range (50°N–30°S)covered during the Atlantic Meridional Transect (AMT) 11 cruise.Five different oceanic provinces were traversed in this cruise:North Atlantic Drift (NADR), North Atlantic Subtropical Gyral(NAST), Canary Coastal (CNRY), Eastern Tropical Atlantic (ETRA),and South Atlantic Gyral (SATL). CNRY and ETRA were affectedby the upwelling Mauritanian and equatorial respectively andprimary production in these provinces was higher than in theoligotrophic subtropical gyres (NAST and SATL). Both mesozooplanktonand phytoplankton biomass were highest around the equator. Theamount of chlorophyll a ingested daily by copepods was noticeablyhigher in mesotrophic than in oligotrophic provinces as shownby the spatial distribution of gut content values and the highabundances of copepods recorded at the equator. Grazing impactalong the transect ranged from 0.2 to 5.6% of the phytoplanktonstanding stock and from 1.6 to 14.5% of primary production.If only phytoplankton >2 µm are considered, the rangesare 1.0–19.4% (stock) and 3.4–44.7% (primary production).Grazing impact upon both phytoplankton biomass and primary productionfollowed a spatial distribution similar to that of chlorophylla ingestion, with higher values in upwelling zones than in thegyres. Weight-specific rates of respiration and NH₄⁺ and PO₄^3–excretion showed large variability both along the transect andwithin provinces, but did not differ between provinces. Therefore,zooplankton assemblages inhabiting the different provinces visitedin the AMT 11 seem to be adapted to the prevailing thermal conditions.Given the substantial proportion of nitrogen and phosphorusthat are supplied to primary producers through the excretoryactivity of mesozooplankton (the percentage of nitrogen andphosphorus requirements of phytoplankton accounted for by mesozooplanktonexcretion was >30% in all the provinces) it follows thatthey may play a crucial role as nutrient regenerators, especiallyin the oligotrophic gyres where regenerated production dominates.     

Macrozooplankton and the persistence of the deep chlorophyll maximum in a stratified lake

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