In situ diel variation in gut pigment contents of Ceriodaphnia sp. in stratification and destratification periods

The short-term, in situ diel grazing of Ceriodaphnia sp. duringperiods of stratification and mixing was investigated usingthe technique of fluorimetric analysis of the gut pigments.There were considerable seasonal differences in feeding behaviourIn mixing, when the concentration of chlorophyll a in the watercolumn was high and Ceriodaphnia abundance was low, gut pigmentcontents showed clear diel variation patterns, probably dueto diel variations of the high values of feeding activity observedin the 24 hour cycle The maximum values were found at dawn.On the other hand, no diel variations in gut pigment were observedduring periods of stratification and while the amounts of pigmentsin the water and in the gut were very low, species abundancewas high. Taking into account the ambient conditions, the authorsdiscuss the possibility that the change of feeding of the Ceriodaphniasp. observed when the environment changed from a mixing periodto one of stratification represents a change from herbivorousto detritivorous behavior.     

A study of time-dependent primary productivity in a natural upper-ocean mixed layer using a biophysical model

A one-dimensional biophysical model of time-dependent photosynthesisin the upper-ocean mixed layer was applied near 36.5°N,74.5°W. The photosynthesis submodel was formulated and parameterizedbased on daylong, light-manipulation experiments performed ontwo phytoplankton communities collected prior to dawn on October13, 1992: one from 5 m depth in the upper mixed layer and theother from 28 m depth below the pycnocline. Time course biologicalmeasurements included chlorophyll a concentration, primary productivityand DCMU fluorescence ratio. The biophysical model was thenused to predict the physical and biological response of thewater column on October 14, 1992. A pre-dawn conductivity-temperature-depth-fluorescence-transmission-rosette(CTDFT-rosette) cast provided data to initialize the water columnstratification and the distribution of phytoplankton biomass.The biophysical model was forced using meteorological and oceanographicmeasurements collected continuously throughout the subsequentdaylight period. CTDFT-rosette casts and measurements of chlorophylla concentration, primary productivity and DCMU fluorescenceratio throughout that daylight period provided the data forcomparison with the model predictions. In general, the biophysicalmodel predicted the physical and biological evolution of thesampled water column. Although the inclusion of vertical mixinginitially improved the accuracy of prediction, agreement decreasedwith time, especially in the lower part of the water column.The one-dimensional model suffered from the effects of excludedhorizontal gradients.     

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     

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.     

Short-term feeding rates of A carrier grani in natural conditions: diurnal variation

Short-term in situ diel grazing activity of adult male and femalesofAcania graniSars. were investigated at a coastal station inthe Alboran sea (SE of Spain). Gut pigment contents were measuredfluorometrically, and the ingestion rates, filtration ratesand daily ratios were calculated from gut fullness, evacuationrates and pigment concentrations in sea water. The results indicatethat there were size-related differences in the amount of foodingested by males and females. Gut contents of females wereon average >1.5 times that for males. Body-volume-specificfeedingestimates were, however, the same for both sexes. Gutpassagetimes, estimated from animals collected at the time of day whengut pigments were maximal, were shorter in females. Both sexesshowed diel changes in feeding activity and faecal pellet productionof about one order of magnitude;the maximum rates were observedat dusk, the minimum rates at midday. There was a directrelationshipbetween copepod grazing and pigment concentration in sea waterwhen a time lag was considered. Daytime and night-time in situincubations using natural sea water enriched withlaboratorycultured algae provided evidence of non-continuous feeding andof light intensity control of the diel patterns. The possibleadaptive advantages of differences in sexual size on feedingbehaviour and diurnal variability are discussed.     

Zooplankton community grazing in a hypertrophic lake (Hartbeespoort Dam, South Africa)

Seasonal abundance as total biomass and specific densities ofthe main herbivorous zooplankton (>60 µm) in hypertrophicHartbeespoort Dam from 1981 to 1983 are described. After springzooplankton biomass maxima each year there followed a mid-summerdecline in the Daphnia population and a shift to a smaller bodiedcladoceran community dominated by Ceriodaphnia concomitant witha change from largely edible phytoplankton species to abundantcolonial Microcystis. In situ community grazing rates were measuredthroughout 1983 using a ¹⁴C-labelled unicellular alga. Integratedcommunity grazing rates measured in the aerobic water columnwere highest in December (260.2%/day) when Daphnia was abundantand the edible component of the phytoplankton was diminishing.Lowest integrated community grazing rates occurred in January–February(19.8–35.3%/day) and July (28.3%/day) when the phytoplanktonwas composed almost entirely of Microcysris, and Ceriodaphniadominated the zooplankton community whilst food availabilitywas low. Feeding in Ceriodaphnia was not hindered by abundantlarge Microcysris colonies; total biomass specific grazing ratewas high when Ceriodaphnia dominated and low when Daphnia dominatedthe zoo-plankton community. Results indicate that in hypertrophicconditions it is unlikely that large filter-feeders such asDaphnia are able to retard or limit the development of Cyanophyceaeblooms by high grazing pressure.     

Pigment gut contents of copepods and deep phytoplankton maximum in the Western Mediterranean

The feeding activity of Centropages typicus in the Western Mediterraneanwas evaluated during the summer stratification period, whena deep phytoplankton and chlorophyll maximum (DCM) develops.The lack of significant day-night differences in gut contentssuggests continuous phytoplankton ingestion, despite nictemeralmigration of this species. Copepods captured in the DCM hadgreater phytoplankton gut contents, both in nocturnal and diurnalsamples. The observed pattern of phytoplankton ingestion andvertical migration implies copepod-mediated upward transportof nutrients, which might contribute to the maintenance of primaryproduction in the photic layers.     

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).     

Temporal variability in copepod fecundity during two different spring bloom periods in coastal waters off Plymouth (SW England)

The temporal variation in egg production of the planktonic copepodsCalanus helgolandicus, Temora longicornis and Pseudocalanuselongatus was studied during two different spring bloom periodsin 1989 and 1990 by weekly sampling at two permanent stationsin coastal waters off Plymouth (SW England). Copepod egg productionwas estimated in situ by incubating individual adult femalesin filtered seawater for 24 h (72 h until hatching for P.elongatus)at the field surface temperature. The relationship between copepodegg production rates and: (i) chlorophyll a concentration (totaland >10 µm size fraction), (ii) temperature and (iii)initial copepod gut pigment content was investigated. The springbloom periods were very different in both years, with the occurrenceof a Phaeocystis sp. bloom in 1990, which negatively affectedthe feeding and fecundity of copepods. Egg production ratesin spring 1989 were significantly correlated with chlorophylla concentration (particularly with the >10 µm fraction),field temperature and copepod gut pigment contents. In spring1990, egg production rates were also correlated with copepodgut pigment contents, but no significant correlations were obtainedwith temperature or with chlorophyll a concentration, as a consequenceof the lower egg production rates obtained during the Phaeocystissp. dominance period. These results show that food availabilityis the factor which mainly affects the fecundity of neriticcopepods in short time periods.     

Feeding, excretion and egg production by individuals and populations of the marine, planktonic copepods, Acartia spp. and Centropages furcatus

Diel variations in vertical distribution, gut pigment content,ammonium excretion and egg production were investigated foradult females of Acartia erythraea and A.pacifica in the verticallymixed Inland Sea of Japan and Centropages furcatus in the stratified,neritic Gulf of Mexico. Gut pigment content and egg productionrate were maximal at night and ammonium excretion was maximalduring the daytime. Neither A.erythraea nor A.pacifica adultfemales showed an apparent diel migration, but the former werehighly concentrated in the surface layer during the afternoon.In contrast, C.furcatus adult females showed a clear diel migration,residing immediately above the bottom during the daytime andbeing concentrated between 10 and 25 m depth during the nighttime.Individual-based data on gut content and excretion and egg productionrates were combined with vertical-distribution data to calculatepopulation values. In the Inland Sea of Japan, the resultantpattern for Acartia spp. reflected the diel variation in physiologicalrates and even distribution of adult females, except for theafternoon, surface aggregation of A.erythraea. In the Gulf ofMexico, the pattern for C.furcatus reflected largely the dielvariation in each rate process and the heterogeneous distributionof adult females in the water column. Elevated nocturnal feedingactivity of these copepods may be due to an endogenous rhythm.The daytime maximum in ammonium excretion and night-time maximumin egg production rate indicated approximate half-day and daytime lags, respectively, after the intake of food until itsconversion into dissolved excreta and released eggs.     

Gut evacuation rates and ingestion rates of Eudiaptomus graciloides measured by means of the gut fluorescence method

Diel variation in feeding rates of Eudiaptomus graciloides wasmeasured in situ with the gut fluorescence method during Mayand June in the eutrophic Frederiksborg Slotss Pronounced dielperiodicity occurred in both periods with highest feeding ratesduring the night. The food concentration as well as the periodicitywas almost constant with depth, indicating that the feedingactivity represents a behavioral periodicity. The gut evacuationmeasured in the laboratory on field-collected copepods was exponentialand temperature-dependent. The exponential decline in gut contentwas found for both copepods feeding continuously and when feedinghad ceased. Considering spatial and temporal variation in feedingactivity, daily ratios were 61–68 body carbon per day,and higher than if calculated from existing laboratory dataof short-term feeding dates.     

Gut pigment destruction by the copepod Acartia clausi

The extent of chlorophyll (Chi) a degradation into colorlessproducts by the copepod Acartia clausi was determined by measuringgut fluorescence prior to fecal pellet production. The efficiencyof pigment degradation of animals fed at different concentrationsof Thalassiosira weiss-fiogii varied between 30.56 and 94.05%,and the quantity of pigments lost during gut passage was directlyrelated to total ingestion. The role of feeding history in pigmentdestruction was tested. The fraction of ingested Chi a degradedby copepods acclimated to a high food concentration (15 ng Chla ml^–1) was greater than the fraction degraded by thoseacclimated to a low food concentration (1.5 ng Chl a ml^–1).The percentage of ingested Chl a that A.clausi transformed intofluorometrically undetected compounds was not constant. At present,knowledge of the pigment destruction process indicates thatChi a and Chi --derived pigments are not considered useful quantitativetracers of the feeding activity of copepods. The results ofthis study suggest a reassessment of the application of thegut fluorescence method to evaluate grazing activity of thecopepod A.clausi     

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.     

Zooplankton-phytoplankton interactions in a eutrophic lake

Enclosure experiments were made in a cyanobacteria dominatedlake (Lake Rotongaio) to assess the impact of zooplankton (>150µm) grazing on algal growth rates and determine the effectof diel and vertical changes in zooplankton grazing intensityand nutrient (NH₄-N) regeneration upon abundance of phytoplankton.The filamentous cyanobacterium Anabaena minutissima var. attenuataand diatom Cyclotella meneghiniana showed a negative linearchange in abundance with a gradient in zooplankton grazing intensity.Phytoflagellates were not grazed and showed a positive linearchange in abundance with increasing zooplankton biomass. Theseeffects, as well as shortening of filament length of Anabaena,were caused by raptorial feeding by the alanoid copepod Boeckellapropinqua which dominated the zooplankton. Phytoplankton growthwas not stimulated by addition of nutrients, suggesting nutrientregeneration was not important. Diel and vertical changes infeeding and NH₄-N regeneration rates were measured in Marchand June 1988. Diel differences were more pronounced in Marchwhen the water column was stratified. Specific feeding rateswere more important than vertical changes in zooplankton biomassin determining community grazing rates in March, but in Junewhen the water column was mixed, vertical distribution of zooplanktonbiomass was important. Zooplankton grazing was an importantloss process for phytoplankton in the lower part of the epilimnionin Lake Rotongaio.     

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.     

Life table responses and population dynamics of four cladoceran zooplankton during a reservoir flood

The life table responses and population dynamics of four cladocerans(Ceriodaphnia lacustris, Daphnia parvula, Diaphonosoma leuchtenbergianumand Moina micrura) were measured during a flood of a large turbidreservoir (Lake Texoma, Oklahoma-Texas). Mean recruitment andlongevity of Ceriodaphnia and Daphnia in life table experimentswere reduced during the flood when abundant silt particles andpicoplankton (<2 µm) replaced algal particles in thewater column. Individual growth rates measured in the life tableexperiments were slower for Ceriodaphnia during the flood. Fieldcollections revealed that Ceriodaphnia and Daphnia populationsdeclined immediately after the arrival of turbid water, andremaining females carried fewer eggs. Populations of Moina andDiaphanosoma increased during the flood, and life table experimentsshowed that these species were able to grow well in silt-ladenwater.     

Zooplankton biomass distribution patterns along the western Antarctic Peninsula (December 2002)

The phytoplankton [chlorophyll a (Chl a)], microzooplankton,mesozooplankton and macrozooplankton biomass and distributionwere studied as part of a multidisciplinary project (Tempano)along the Antarctic Peninsula during December 2002. Even thoughthe summer phytoplankton bloom was not yet developed in thearea, autotrophs dominated the plankton biomass. Phytoplanktonvertical distribution was, in general, homogeneous in the upper40–50 m of the water column, further decreasing with depth.Protozoans showed low biomass; their contribution to the totalplankton being one order of magnitude lower than that of autotrophs.The vertical distribution of protozoans was variable among stationswith marked peaks at depths ranging from 30 to 80 m. Mesozooplankton-integratedbiomass was generally low, although there was a notable increasesouthward near the ice marginal zone. Macrozooplankton distributionwas more variable without any clear zonal distribution pattern.The vertical distribution of meso- and macrozooplankton (>4mm) biomass showed clear peaks of abundance comprising differentspecies depending on the geographical area. Our biomass distributiondata suggest a food-web scenario in which macrozooplankton arepreying on mesozooplankton populations only in the northernerstations, and mesozooplankton are, in their turn, shaping theabundance of the emerging populations of microzooplankton. Phytoplankton,on the other hand, seem to be hardly controlled by grazing activity.     

The presence of chlorophyll b and the estimation of phaeopigments in marine phytoplankton

A reverse-phase h.p.l.c. technique was used to estimate theconcentration of chlorophyll b in phytoplankton cultures, fecalpellets of Calanus pacificus, and suspended paniculate matterfrom the Central North Pacific, Oregon coastal waters, and DabobBay (a temperate fjord in Puget Sound, WA, USA). The purposewas to assess the distribution of this pigment in the euphoticzone and its effect on the fluorometnc estimation of phaeopigments.Analyses of natural waters confirm high chlorophyll b concentrations(median mass ratio of b:a > 0.3) at the depth of the chlorophylla maximum in tropical waters while values for temperate planktonare relatively low (median mass ratio of chl b:a = 0.05) andpatchy. Zooplankton fecal pellets showed a significant enrichmentin chlorophyll b, suggesting grazing as a mechanism to explainhigh concentrations of this pigment at the bottom of the euphoticzone. It is estimated that the presence of chlorophyll b couldcause an average overestimation of phaeopigment concentrationby the fluorometnc technique of 38% between 0 and 200 m in theCentral North Pacific. This effect is more pronounced at thelayer of chlorophyll b maximum (120–140 m). ¹Present address: Marine Biology Research Division, A-002, ScrippsInstitution of Oceanography, La Jolla, CA 92093, USA     

Interrelations between the vertical distribution of Daphnia and chlorophyll a in two large limnetic enclosures

This paper summarises the major factors influencing the daytimevertical distribution of Daphnia hyalina var. laciutris (Sars)in two large experimental enclosures (Lund Tubes). In both tubes,the depth at which the Daphnia aggregated during the day wasclosely related to sub-surface ir-radiance. On a few occasionsaggregations of Daphnia were found in ‘dark’ waterassociated with a deep chlorophyll maximum. On many occasions,however, the animals' light response per se ensured their aggregationat depths of maximum phytoplankton abundance. The most importantfactor influencing the dispersion of animals in the water columnwas the steepness of the light gradient. In turbid water verticalaggregations were well defined, whereas in clear water the animalswere widely dispersed around their depth of maximum abundance.Daphnia also tended to disperse in the water column when theirpopulation density was high or when food was scarce. A simplemodel based on surface irradiance, water turbidity and populationdensity explained the basic pattern of vertical distributionthroughout the season. The implications of these findings arediscussed in relation to the effects of zooplankton grazingand speculations on the adaptive significance of depth regulation.     

HPLC analysis of phytoplankton pigments from Lake Kinneret with special reference to the bloom-forming dinoflagellate Peridinium gatunense (Dinophyceae) and chlorophyll degradatio products

Photosynthetic pigments extracted from the paniculate materialof the water column of Lake Kinneret were studied throughoutthe periods of May 1988-June 1989, and November 1993-November1994, by means of HPLC. The temporal and vertical variationof the pigment suite found agreed with the microscopically determinedphytoplankton record. The regression calculations of taxon-specificbiomass with the corresponding signature pigments suggest thatpigment analysis may be a useful tool for the monitoring ofbloom-forming species, e.g. the dinoflagellate Peridinium gatunenseNygaard. The HPLC pigment analysis permitted the identificationand quantification of chlorophyll degradation products, providingfor the first time information about their composition in LakeKinneret. Chlorophyllide a was the major detectable degradationproduct of chlorophyll a, varying between 1 and 9% of the chlorophylla concentration. Other chlorophyll a derivatives appeared mostlyin minor quantities. Pheophytin a was virtually lacking in allthe samples. Removal rates of pigments, measured by sedimentationtraps, indicated that the degradation of chlorophyll a via chlorophyllidea is a dynamic process that continues during the sedimentationof the phytoplankton particles.