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.     

The fate of the carotenoid pigment fucoxanthin during passage through the copepod gut: pigment recovery as a function of copepod species, season and food concentration

The amount of fucoxanthin, a taxonomically diagnostic carotenoid,recovered after passage through the guts of the copepods Acartiacalifomiensis and Calanus pacificus, was determined after thecopepods had fed on low (50 µg Cl^–1) and high (350µg C1^–1 for Acartia; 500 ug C H for Calanus) concentrationsof the diatom Thalassiosira weissftogii, during spring (May)and winter (December). Changes in pigment concentrations andcell abundances were assessed in experimental (with copepods)and control (without copepods) samples by standard incubationexperiments. Pigment recovery was assessed by (i) comparingthe amount of ingested pigment recovered in the experimentalgroups with that predicted to have been ingested from cell countdata and (ii) comparing fuco-xanthin/cell ratios in controland experimental samples. Both techniques suggested that pigmentloss is substantial (usually 60–100%), regardless of species,food availability or season. Patterns of pigment conservationdiffered between species, although pigment recovery was alwayshigher at high, than at low, food concentrations. Pigment recoveryin Acartia was higher (9.4–28.0%) in the spring than duringthe winter (0 recovery), regardless of food concentration. InCalanus, however, pigment recovery was always higher at high(34.9–67.8%) than at low (0 recovery) food concentrations,regardless of season.     

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     

Developmental rates of the copepods Calanus australisand Calanoides carinatus in the laboratory, with discussion of methods used for calculation of development time

The copepods Calanus australis and Calanoides carinatus developedfrom egg to adult in 20.3 and 18.3 days respectively at 15.5?Cand 16.0 and 12.0 days at 19.5?C. For both species the durationof the first two naupliar stages was short, <1 day. The thirdnaupliar stage was prolonged in C.australis but short in C.carinatus. Isochronal development was approximated from fourthnauplius through third copepodite; fourth and fifth copepoditestages were relatively long. Only female C.australis were produced;60% of the C.carinatus were female. Equiproportional developmentwas found for C.carinatus but not for C.australis. For bothspecies the variance in stage duration was established at thethird nauplius then remained unchanged through the older stages.In calculating developmental rates several commonly used methodswere compared. The method of ‘first appearance of an individualof a given stage’ and ‘mean time of appearance ofa stage’ both yielded estimates of development time thatwere 10–15% faster than the more commonly used methodof ‘median development time’. We suggest that allfuture work on developmental rates of copepods should use astandardized method for the calculation of stage-specific developmentalrates and that ‘median development time’ be themethod of choice. ³Present address: Marine Sciences Research Center, SUNY, StonyBrook, NY 11794-5000, USA     

Gut clearance and pigment destruction in a herbivorous copepod, Acartia tonsa, and the determination of in situ grazing rates

Gut clearance rates of starving and continuously feeding Acartiatonsa were estimated. During the initial 30 min the rates weresimilar (0.045 and 0.048 min^–1, respectively; 14°C)but thereafter starving animals expelled the remains of theirgut contents at half the rate (0.019 min^–1) of fed ones(0.048). Pigment destruction was estimated by (i) incubationexperiments over 3–4 days, (ii) silica to pigment ratioin algae and faeces and (iii) by gut filling experiments. Theincubations showed that 8% of the ingested pigments were destroyedto nonfluorescent residues during gut passage. The silica topigment ratio method gave an average of 11 % (1 –24) destructionand gut-filling experiments showed no systematic differencebetween ingestion measured as gut filling rate (fluorescence)and particle reduction. ¹Present address: Kristineberg Marine Biological Station, S-45034 Fiskebäckskil, Sweden     

Egg production and hatching success in the calanoid copepods Calanus helgolandicus and Calanus finmarchicus in the North Sea from March to September 2001

Spatial and seasonal egg production rates (E_r) and egg hatchingsuccess in the copepods Calanus finmarchicus and Calanus helgolandicuswere measured in the North Sea from March to September. Foodavailability was monitored by chlorophyll and protist concentrationsand three size fractions of seston fatty acids. Seasonal andspatial distribution and production differed between the species.Calanus finmarchicus was found only offshore of the 50-m isobath,with decreasing E_r (37–28 eggs female^–1 day^–1)from March to July. Calanus helgolandicus had two abundancepeaks, in spring and autumn, with a low in May during whichtime the highest E_r were observed (38 eggs female^–1 day^–1).At other times, E_r in C. helgolandicus remained lower than inC. finmarchicus (     

The relative importance of food and temperature to copepod egg production and somatic growth in the southern Benguela upwelling system

The fecundity and somatic growth rates of Calanus agulhensisand Calanoides carinatus, the dominant large calanoid copepodsin the southern Benguela upwelling system, as well as the fecundityof several other common copepods, were measured between Septemberand March of 1993/94 and 1994/95. Mean egg production of mostcopepods was low at >30 eggs female^-1 day^-1 {Calanoides carinatus23.7, Calanus agulhensis 19.0, Neocalanus tonsus 16.1 and Rhincalanusnasutus 26.1), whereas the mean fecundity of Centropages brachiatuswas significantly greater (83.6 eggs female^–1 day^-1).This study also presents the first comprehensive field estimatesof the fecundity of Nanno-calanus minor (mean: 26.1 eggs female^–1day^–1, range: 0.0–96.2 eggs female^–1 day^–1)and of somatic growth of N6 and all copepodite stages of Calanoidescarinatus (decreasing from 0.58 day^–1 for N6 to 0.04 day^–1for C5). Somatic growth rates of Calanus agulhensis also declinedwith age: from 0.57 day1 for N6 to 0.09 day1 for C5. Data ongrowth rates were used to assess the relative importance offood [as measured by total chlorophyll (Chi) a concentration],phytoplankton cell size (proportion of cells >10 µm)and temperature to the growth of copepods. Multiple regressionresults suggested that fecundity and somatic growth rates werepositively related to both Chi a concentration and phytoplanktoncell size, but not to temperature. Although it was not possibleto separate the effects of Chi a concentration and phytoplanktoncell size, data from previous laboratory experiments suggestthat copepod growth is not limited by small cells per se, butby the low Chi a concentrations that are associated with theseparticles in the field. Despite growth not being directly relatedto temperature, a dome-shaped relationship was evident in somespecies, with slower growth rates at cool (<13°C) andwarm (>18°C) temperatures. The shape of this relationshipmirrors that of Chi a versus temperature, where poor Chi a concentrationsare associated with cool and warm temperatures. It is concludedthat the effect of food limitation on growth of copepods outweighsthat of temperature in the southern Benguela region. Sourcesof variability in relationships between growth and Chi a concentrationare discussed.     

Feeding and metabolism of the siphonophore Sphaeronectes gracilis

The in situ predation rate of the siphonophore Sphaeronectesgracilis was estimated from gut content analysis of hand-collectedsiphonophores and from laboratory data on digestion rates ofprey organisms. At daytime prey densities of 0.25 copepods 1^–1,S. gracilis was estimated to consume 8.1 – 15.4 prey day^–1siphonophore^–1. From data on abundances of siphonophoresand copepods, S. gracilis was estimated to consume 2–4%of the copepods daily. In laboratory experiments, ingestionrates averaged 13.8 prey day^–1 siphonophore^–1 atprey densities of 5 copepods 1^–1 and 36.9 at 20 copeods1^–1. This was equivalent to a specific ingestion rate(for both carbon and nitrogen) of –17% day^–1 and45% day^–1, respectively, while specific ingestion in situwas only 2% day^–1. Ammonium excretion averaged 0.095 µg-atsiphonophore^–1 day^–1 at 5 prey 1^–1, and 0.162at 20 prey 1^–1. The specific respiration (carbon) andspecific excretion (nitrogen as ammonium) were calculated tobe 3% day^–1 at the lower experimental food level, and5% day^–1 at the higher food level. ¹Contribution from the Catalina Marine Science Center No. 66. ²Present address: Dept. of Biology, University of Victoria,Victoria, B.C., Canada V8W 2Y2.     

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.     

Egg production and hatching success in the peri-Antarctic copepod Calanus simillimus

In situ rates of egg production and hatching success are reportedfor Calanus simillimus, one of the most abundant calanoid speciesin peri-Antarctic regions, during the Italian ‘Italics’cruise in the Straits of Magellan in March-April 1995. Low fecundity(8.6 eggs female^–1 day^–1) and fecal pellet production(0.8 fecal pellets female^–1 day^–1) in this periodindicate that the species was feeding very poorly. Sixty-sixof the 126 females sampled did not produce eggs and 80 of thesedid not produce any fecal pellets during the 24 h period ofincubation. Striking abnormal naupliar and embryonic developmentwas recorded in 81.8% of the eggs spawned. Aberrant eggs didnot undergo normal cleavage, and failed to develop to hatching.Deformed nauplii were asymmetrical and presented strong anatomicalanomalies in the total body length and number of swimming appendages.These results are discussed in the light of recent findingson the causes of low hatching success in copepods.     

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.     

The energetics of Calanus euxinus: locomotion, filtration of food and specific dynamic action

The effect of locomotor activity on respiration rate was studiedin the food-deprived copepod Calanus euxinus tethered to a forcesensor. The power generated by mouth appendages during cruisinglocomotion, with a frequency of 40 Hz, accounted for 0.026 and0.0031 W for metabolic and mechanical processes, respectively.To overcome total hydrodynamic drag during foraging with a meanswimming speed of 3.2 cm s^–1, the copepods need 0.4 x10^–3 W, equating to 1.3% of total metabolism. The lossesof mechanical energy for body propulsion amounted to 1.3 x 10^–3W, whilst the cost of feeding current generation run up to 1.8x 10^–3 W, or 58% of the total. Changing of locomotor activityand respiration rate during feeding was examined separatelyin tethered and free-swimming copepods. At algal concentrationof 300 µg C L^–1, the magnitude of specific dynamicaction (SDA) averaged 1.2 ± 0.44 nL O₂ µg C^–1h^–1 in copepodites V and females, with similar movingactivity before and during feeding. The contribution of SDAinto total metabolism varied from 23 to 85% in C. euxinus withlow activity level and constituted only 10% in active animals.In starved copepods, with low locomotor activity, feeding eventsstimulated the increase in frequency and total duration of locomotionwhich resulted in elevated energy expenditure enhancing the‘apparent SDA’.     

Remote detection of algae by copepods: responses to algal size, odors and motility

Selective feeding on large algae by copepods involves remotedetection of individual particles and subsequent active captureresponses In this study we use radiotracer experiments to quantifythe clearance rates of five coexisting freshwater copepods andto investigate the relative merits of the chemoreception andmechanoreception hypotheses of remote detection Tropocyclopsand three diaptomid copepods exhibited relatively high clearancerates when feeding on low concentrations of large algae, suggestingthat most previous studies with freshwater copepods have underestimatedmaximal clearance rates and the degree of size selectivity.All five species of copepods exihibited strong selection foran intermediate-sized flagellate (25 µm Cartena) or alarge-sized nonmotile alga (80 µ.m Pediastrum) over asmall-sized flagellate (6 µ.m Chlamydomonas). The weight-specificclearance rate for Tropocylops prasmus feeding on motile Cartena(271 ml mg^–1 h^–1) was about twice that of threediaptomid copepods and more than an order-of-magnitude higherthan the estimate for Epischura lacustris feeding on its preferredalga, Pediastrum Assuming that distance chemoreception (‘smell’)is important in remote detection, we predicted that the additionof high concentrations of ‘algal odors’ would obscureany chemical gradients emanating from individual algal cellsand would thereby hinder the remote detection and active captureof large algae Contrary to this hypothesis, the addition ofamino acids, sucrose, and algal extracts had no effect on theclearance rates and selectivity of Diaptomus birgei. These results,together with recent cinematographic studies (Vanderploeg etal.,1990), suggest that mechanoreception is the primary mechanismfor the remote detection of large particles by diaptomid copepods.A raptorial cyclopoid, Tropocyclops prasinus, exhibited strongpreferences for motile algae, whereas a suspension-feeding calanoid,D birgei, did not select between motile and nonmotile cells.Motility appears to be an important factor in algal detectionfor small cyclopoid copepods but not for suspension-feedingdiaptomids     

Autotrophic and heterotrophic nanoplankton in the diet of the estuarine copepods Eurytemora affinis and Acartia bifilosa

The ingestion of autotrophic and heterotrophic nanoplanktonby two estuarine copepods, Eurytenora affinis and Acarith bifilosa,was measured in various environmental conditions using the incubationmethod and epifluorescence microscopy. Egg production of thespecies was also deter mined in order to estimate their carbonrequirements. Assuming a gross efficiency of egg productionof 0.3, nanoplanktonic carbon ingested always met the carbonrequirements suggesting that, most of the time, other carbonsources could be unnecessaly. Nanoplankton ingestion by A.bifilosa(from 128 to 1693 cells ind.^–1 h^–1) was dominatedby autotrophic forms (60–97%) and was seriously affectedby high (>100 mg l^–1 suspended particulate matter (SPM)concentrations. Nanoplankton ingestion by E.affinis (from 300to 1049 cells ind^–1 h^–1) was relatively stable incomparison, but this latter species seemed to switch its grazingpressure from autotrophic to heterotrophic forms when SPM concentrationsincreased. Thus, two copepod species, living in the same estuary,presented two different feeding behaviours, probably to maximizeenergy input per unit of energy expenditure. Such differencescould contribute to the spatial and seasonal segregation ofthese species which is usually observed.      

Rates of ingestion and their variability between individual calanoid copepods: direct observations

The goals of this study were to determine rates of ingestionand fecal pellet release, and their variability, for individualplanktonic copepods over extended periods of time (>20 min).Ingestions and rejections of individual cells of the diatomThalassiosira eccentrica by adult females of the calanoid Paracalanusaculeatus were directly quantified by observing individual copepodscontinuously at cell concentrations ranging from 0.1 toli mm³l^–1.Average ingestion rates increased with increasing food concentration,but were not significantly different between 0.3 and 1.0 mm³l^–1(9.8 and 32.7 µg Cl^–1) of T.eccentrica. Rates ofcell rejections were low and similar at 0.1 and 0.3, but weresignificantly higher at 1.0 mm³ l^–1. The coefficientsof variation for average ingestion rates of individual copepodshardly differed between food concentrations, ranging from 17to 22%, and were close to those for average fecal pellet releaseintervals which ranged from 15 to 21%. A comparison betweenindividuals at each food concentration found no significantdifferences at 1.0; at 0.1 and 0.3 mm³l^–1, respectively,ingestion rates of four out of five females did not differ significantlyfrom each other. Average intervals between fecal pellet releaseswere similar at 0.3 and 1.0 mm³l^–1 of T.eccentrica, butsignificantly longer at 0.1 mm³ l^–1. Fecal pellet releaseintervals between individuals were significantly different ateach food concentration; these significant differences wereattributed to rather narrow ranges of pellet release intervalsof each individual female. Potential sources/causes of variabilityin the sizes and rates of copepods in the ocean are evaluated. ³Present address: Great Lakes Environmental Research Laboratory,2205 Commonwealth Boulevard, Ann Arbor, Ml 48105, USA     

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.     

Effects of prey motility and concentration on feeding in Acartia tonsa and Temora longicornis: the importance of feeding modes

Feeding experiments were conducted with the ambush-feeding copepodAcartia tonsa and the feeding-current-generating copepod Temoralongicornis. The copepods were offered a mixed diet of the dinoflagellateHeterocapsa triquetra and the ciliate Balanion comatum of similarcell size. The dinoflagellate was offered at a constant concentrationof 10–15 cells mL^–1, whereas the ciliate was offeredat a variety of concentrations, ranging from 7 to 57 cells mL^–1.Copepods with different feeding modes possess different mechanismsfor prey detection, suggesting that the two copepods would responddifferently to the two prey types. Both copepods had significantlyhigher clearance rates on the highly motile ciliate than onthe less motile dinoflagellate. In encounters between A. tonsaand its prey, we argue that this is due to the higher hydromechanicalsignal generated by the ciliate. The advection feeding copepodT. longicornis fed on the two prey according to their relativeconcentrations; in this case, we suggest that although B. comatumis capable of detecting feeding-current-generating predators,the feeding current velocity generated by T. longicornis isgreater than the escape velocity of this ciliate.     

Population biomass, feeding, respiration and growth rates, and carbon budget of the scyphomedusa Aurelia aurita in the Inland Sea of Japan

We investigated the seasonal occurrence, wet : dry : carbon: nitrogen weight ratios, population biomass, gastric pouchcontents, and rates of feeding, growth and respiration of thescyphomedusa Aurelia aurita in the central part of the InlandSea of Japan. Aurelia aurita medusae began to appear in January/Februaryas ephyrae, reached annual maximum body size in July/August,and disappeared, presumably due to death, by November. Initialslow growth in early spring was followed by a period of exponentialgrowth (mean growth rate: 0.069 d^–1) between April andJuly. In the Ondo Strait, which is characterized by strong tidalmixing, the A. aurita population (mean carbon biomass: 66.0mg C m^–3) overwhelmingly dominated the zooplankton-communitybiomass (mean biomass of micro- and mesozooplankton: 23.7 mgC m^–3) between May and early August The gastric contentanalysis revealed that A. aurita ate almost all micro- and mesozooplankters,of which small copepods were most important. On the basis ofdigestion time for small copepods (60 min) and their abundancein the gastric pouch of field-collected A. aurita, we determinedthe weight specific feeding rates and clearance rates. The formerincreases linearly with increasing copepod abundance, but thelatter was relatively constant irrespective of the food supply.We also measured the respiration rates of A. aurita and expressedthem as functions of body weight and temperature. These physio-ecologicalparameters enabled us to construct the carbon budget of theA. aurita population typical of early summer in the Ondo Strait.Predicted population-feeding rate (6.07 mg C m^–3 d^–1)was higher than the population-food requirement for both metabolismand growth (4.55 mg C m^–3 d^–1), indicating thatfood supply was sufficient to sustain the observed growth rate.This feeding rate was equivalent to 26% of micro- and mesozooplanktonbiomass, a significant impact on zooplankton.     

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.     

Feeding, growth and food conversion of the marine cladoceran Penilia avirostris

Ingestion and clearance rates, feeding behaviors and life historyvariables of the marine cladoceran Penilia avirostris were evaluatedover a range of food concentrations encountered in nature (0.01–3.0mm³ 1^–1 of Isochrysis galbana). Ingestion rates increasedand clearance rates decreased with increasing food concentration.No maximum feeding thresholds were observed over the range ofalgal concentrations offered. Weight-specific ingestion ratesdecreased with increasing body weight. Feeding behaviors suchas mandibular activity, abdominal claw rejections of cloggedfeeding structures and feeding appendage activity decreasedat a food level of 0.3 mm³ l^–1 of l. galbana. Peniliaavirostris had very poor survivorship at extremely low (0.01mm³ l^–1 and high (3.00 mm³ l^–1) food levels. Mortalitywas hardly affected at food levels of 0.03–1.0 mm³ l^–1Reproduction did not occur at food levels of