Feeding of freshwater filter-feeders at very low food concentrations: Poor evidence for "threshold feeding" and "optimal foraging" in Daphnia longispina and Eudiaptomus gracilis

The feeding behaviour of two freshwater zooplankton species,the calanoid copepod Eudiaptomus gracilis and the cladoceranDaphnia longispina, at extremely low food concentrations hasbeen studied in order to test whether the animals react as predictedby the models of optimal foraging. Three different sized algae were offered in concentrations downto 1µg C/litre at 7°C during winter and 19°C duringsummer. Ingestion rates were measured by use of a flow-through¹⁴C-technique In contrast to the findings of several authors working withmarine copepods a threshold concentration where the animalsceased filtering could be detected only in one experiment byregression analysis (Daphnia, 19°C, food: Stichococcus).A depression of the filtering rate at low concentrations wasobserved only in Daphnia at 19°C with Stichococcus and Scenedesmusas food. It is supposed that in this case the reduction of thefiltering rate is not a direct response of the filtrator, butis caused by exhaustion due to the experimental conditions. As freshwater zooplankton is very rarely exposed to food abundancesas low as in the marine environment, there seems to be no selectionpressure favouring the evolution of an "optimal forager" behaviour. ⁽¹⁾Supported by Deutsche Forschungsgemeinschaft     

Demography of a natural population of Daphnia retrocurva in a lake with low food quality

The hypothesis was tested that Daphnia exhibit low populationrecruitment in a lake with phosphorus (P)-deficient food, whereDaphnia populations failed to increase in response to greatlylowered minnow density. Egg ratio analysis was performed onDaphnia retrocurva from L110 of the Experimental Lakes Area,Ontario, in 1994. L110 has high food abundance (seston carbon(C) 100 µM C), but the seston is P deficient (C:P atomic- 600). Chrysophytes dominated the algal community. The fishcommunity in this lake was manipulated in 1993 and 1994 by theaddition of northern pike, Esox lucius. Zooplanktivorous minnowswere greatly reduced after the piscivore addition, but the expectedincrease in abundance of the Daphnia to this altered predationregime was not observed, If food quality constrains the responseof the Daphnia to changed predation, there should be low recruitment in the relatively sparse Daphnia population in this lake.Birth rates estimated by standard egg ratio analysis were <0.1day^–1 for most of the sample period. In comparison toannual averages from the literature, recruitment in L110 wasmoderately, but not dramatically, low. Daphnia exhibited pronounced‘normal’ vertical migration (up at night, down duringthe day) all year, and a strongly declining size at first reproductionthrough the season. Population density and birth rates seemedto show much less short-term change than previously observedin other lakes. Although the demo graphic predictions for foodquality constraints seemed clear, in situ demography unfortunatelyneither supported nor strongly rejected the overall hypothesisof food quality limitation in this lake with very high C:P ratios.Possibly, direct demographic effects of low food quality willbe difficult to observe in nature.     

Resource depression in Daphnia galeata, Daphnia cucullata and their interspecific hybrid: life history consequences

In this study, the effect of the food concentration on severallife history parameters of Daphnia galeata, Daphnia cucullataand the hybrid between these two species was examined. The foodconcentration had a significant effect on both growth and reproductionof all three taxa. Mortality was highly species specific, andto a lesser extent dependent on the food concentration. Changesin life history parameters resulted in changes in the intrinsicrate of population increase (r). The food dependence of thedifferent species was found to be different. Daphnia cucullatahad a relatively constant and low r. The r values for the hybridvaried most, with low values compared with the parental specieswhen grown at low food levels and relatively high values athigher food concentrations. Our results suggest that when foodconcentrations are high the hybrid will have a selective advantageover both parental species, but that at lower food levels D.galeatawill have an advantage over both D.cucullata and the hybrid.     

The influence of autotrophic and heterotrophic foods on the demography of Daphnia longispina under starved, semi-natural and enriched conditions

We tested the influence of different types of food on the fitnessof Daphnia longispina during a period of poor resources by comparingthe growth and reproduction of individuals fed on differentdiets. When Daphnia were under starvation conditions, both dissolvedorganic carbon and metabolic products enhanced bacterial growth,and the cladoceran produced 15 neonates and survived for the55 days of experiment. In the natural water, the edible foodconsisted essentially of bacteria and heterotrophic flagellates,which enabled the cladoceran to grow and reproduce. In waterenriched with Cryptomonas (an alga) or Chilomonas (a heterotrophicflagellate), the daphnid growth rates increased considerablyfrom generation to generation, but later in the time courseof the experiment with Chilomonas than with Cryptomonas. Thisalga enhanced the phosphorus:carbon ratio. The primiparous daphnidsfed with Cryptomonas were smaller than those fed with the heterotrophicflagellate and enrichment with the heterotrophic flagellatealso enhanced reproduction. In Lake Tazenat, when the algalbiomass was insufficient to support the Daphnia life cycle,the heterotrophic organisms were an essential food source. Whenfeeding essentially on bacteria, Daphnia has the capacity togrow and reproduce at very low food concentrations, and thisis important in determining its fitness under competitive conditions.     

Food quality of detritus derived from the filamentous cyanobacterium Oscillatoria limneticafor Daphnia galeata

Detritus derived from the filamentous cyanobacterium ,Oscillatorialimnetica was fed to Daphnia galeataDetritus supported growthand reproduction comparable to that on the green alga Scenedesmusobliquus. The live filaments of O.limnetica were, however, oflower food quality. Biochemical parameters of these food types,thought to be important in Daphnia nutrition, were also determined.It is concluded that detritus can be a good quality food forDaphnia, making eutrophic lakes better food environments thangenerally assumed. ¹Present address: Department of Applied Chemistry and Microbiology,Division of Microbiology, BiocenterViikki, FIN-00014 HelsinkiUniversity, Finland     

Diel and vertical variability of seston food quality and quantity in a small subalpine oligomesotrophic lake

Vertical profiles of seston food quality and quantity were measuredin subalpine Castle Lake for particulate carbon, chlorophylla, fatty acids and phosphorus in addition to abiotic parametersincluding water temperature and dissolved oxygen levels. Fieldand laboratory incubation experiments were employed to manipulateDaphnia rosea growth environments. Sestonic eicosapentaenoicacid (EPA) content was much lower, whereas carbon to phosphorus(C:P) ratios were much higher in the epilimnion compared tothe deep-water chlorophyll maximum (16–20 m). In a temperature-controlledincubation, Daphnia grew faster when fed seston from the deep-waterchlorophyll maximum. In in situ cage incubations, D. rosea grewfastest in the epilimnion. Daphnia rosea in a migrating treatmentexhibited intermediate growth rates between the ones for epilimnionand hypolimnion. A projection of D. rosea growth rates by growthmodels without regard to water temperature showed much highergrowth potentials in the hypolimnion. However, with inclusionof water temperature, epilimnetic water always gave higher potentialgrowth rates. In this lake, warmer temperatures of epilimneticwater override the higher food quality and quantity providedby the deep-water chlorophyll maximum.     

Midsummer decline of a Daphnia population attributed in part to cyanobacterial capsule production

A midsummer decline in a Daphnia population has been documentedin Lake Winnebago, Wisconsin, USA, similar to that which typicallyoccurs in other freshwater lakes throughout the world. We investigatedthe role of two products of the cyanobacterium Microcystis aeruginosa(microcystin and polysaccharide capsular matrix) in contributingto this population decrease by utilizing laboratory and lakestudies. Feeding on either the microcystin-producing, minimallyencapsulated M. aeruginosa strain PCC 7820 or the highly encapsulated,non-microcystin-producing strain C3-40, resulted in decreasedsurvival when compared with animals fed chlorophytes. The survivorshipof Daphnia fed C3-40 cells washed to remove the capsule increasedby 30% over that seen in animals fed encapsulated C3-40. Feedingpurified microcystin or capsule alone decreased Daphnia survivalto a greater degree than did starvation. Physiological studiesconducted while Daphnia were exposed to these food sources showedthat ingestion of capsular material resulted in increased post-abdominalclaw movements and decreased mandibular movements. Concurrently,elevated respiration rates were measured in Daphnia, implyingthat capsule increased the energy expended by these animalsthrough increased attempts to reject the material and decreasedfood intake. Lake studies reflected the results of the laboratoryexperiments. The midsummer decline of the Daphnia populationoccurs as the Microcystis biovolume increases and both microcystinand capsular matrix levels rise. While both cyanobacterial productsmay contribute to the midsummer decline in Daphnia pulicaria,laboratory studies suggest that encapsulation may play the greaterrole.     

Is the midsummer decline of Daphnia really induced by age-0 fish predation? Comparison of fish consumption and MDaphnia mortality and life history parameters in a biomanipulated reservoir

We analysed the temporal pattern of a Daphnia galeata populationand the development of the age-0 Ash community in a long-termbiomanipulated lake (Bautzen reservoir, Germany) during springand early summer of two successive years. In Bautzen reservoir,the age-0 Rsh-Daphnia interaction is a key process within thefood web due to the low abundance of adult zooplanktivorousfish. Daily consumption of daphnids by age-0 fish, as estimatedby a bioenergetics model, was compared to daily mortality ratesof daphnids. In addition, we estimated life history parametersof Daphnia that may indicate predation impact by fish. A midsummerdecline of daphnids occurred only in July of 1996, whereas in1995 the Daphnia biomass remained >4 mg wet weight 1^–1for the entire summer. The percentage of total Daphnia mortalitydue to fish predation before the onset of the midsummer declinewas –2% day^–1. Temporal patterns of individual size,clutch size and size at maturity also indicated that the 1996midsummer decline of daphnids was not the exclusive consequenceof age-0 fish predation. Instead, low reproductive capacityof daphnids also contributed significantly to the decline. Consequently,year-to-year variation of the Daphnia dynamics may be determinedby a fine-tuned ‘timing’ between the period of reducedfecundity and the time of the strongest predation impact byage-0 fish.     

Can phaeopigments be used as markers for Daphnia grazing in Lake Constance?

The formation of chlorophyll a degradation products was measuredwith natural phytoplankton from Lake Constance and Daphnia magnaand native Daphnia as grazers in grazing experiments duringspring bloom conditions using high-pressure liquid chromatography(HPLC). Chlorophyll a start concentrations were between 1.2and 16.3 µg l^–1; phaeopigment weights constituted5% of chlorophyll a weight. Only phaeophorbide a was a markerfor Daphnia grazing; concentrations of other phaeopigments (phaeophytina, chlorophyllide a and two unidentified phaeopigments) didnot increase during Daphnia grazing. Conversion efficiencies(chlorophyll a to phaeophorbide a) were between 0 and 43% ona weight basis, and between 0 and 65% on a molar basis. Conversionefficiencies were highest at high grazer density (40 Daphnial^–1) and after a 24 h exposure time. Grazing by microzooplanktonprobably led to the formation of the two unidentified phaeopigments.In Lake Constance, Daphnia density was significantly positivelycorrelated with the phaeophorbide a/chlorophyll a ratio whenit was <5000 Daphnia m^–3. However, when higher Daphniadensities were included in calculations, then Daphnia densitywas positively, but insignificantly, correlated with the phaeophorbidea/chlorophyll a ratio. This suggests that when the level offood per Daphnia is low, then grazing is more efficient withless production of phaeophorbide a and a higher production ofcolourless products.     

Does interference competition from Daphnia affect populations of Keratella cochlearis in Loch Leven, Scotland?

A very marked inverse relationship between Daphnia hyalina var.lacustris Sars and Keratella cochlearis (Gosse) population densitieswas observed in Loch Leven, Scotland, UK between 1978 and 1982.The natural death rates of the rotifer population were far lowerthan would have been expected in response to interference competitionfrom Daphnia. Keratella birth rates fell, along with chlorophylla concentrations, when Daphnia filtration rates were high. Theresults indicate that, when Daphnia were abundant, direct competitionfor food was the most likely factor suppressing Keratella populationgrowth.     

Experimental studies of food and suspended sediment effects on growth and reproduction of six planktonic cladocerans

Life-long somatic growth and egg production responses of Daphniabarbata, Daphnia gibba, Daphnia laevis, Daphnia pulex, Moinamicrura and Diaphanosoma excisum were examined in relation todifferent food and suspended sediment levels, in factorial experimentsat 20°C. Food treatments involved additions of 0, 100, 500and 2500 µg C l^–1 of Selenastrum to 20 µmfiltered water, generally from source lakes. These food levelswere tested at two suspended sediment levels in all species,viz. (i) natural source-lake nephelometric turbidities of 7–65nephelometric turbidity units (NTU) ({small tilde}10–70mg l^–1 TSS) and (ii) {small tilde}3- to 4-fold enrichmentof natural levels (8-fold in D.excisum). A third sediment-freetreatment was tested in all species of Daphnia other than D.gibba.Growth in body length and reproductive output were assesseddaily for individual animals. Main and interactive effects offood and sediment on growth were assessed at selected ages fromday 3 through subsequent life using analysis of variance (ANOVA).Strong and highly significant positive food effects were apparentat all ages in all species examined, with the curious exceptionof D.excisum. The onset of maturity and fecundity were alsopositively related to food level. Sediment effects on growth,age of maturity and fecundity were variable, but tended to bemuch weaker than food effects. Both stimulatory (positive, especiallyat low food levels) and inhibitory (negative) effects of sedimenton growth were apparent, both between and within species. Performancein sediment-free water was very poor, except in D.pulex. Food-sedimentinteraction effects were variable. Generally, turbidwater specieswere influenced less adversely by sediment than clear-waterspecies, indicating the existence of environmentally appropriateadaptive responses. Overall, the findings indicate that lightlimitation of algal resources by suspended sediments, ratherthan the condition of high turbidity per se, is responsiblefor the apparently negative impact of suspended sediments frequentlyobserved on natural populations of cladoceran zooplankters.     

Influence of two Daphnia species on summer phytoplankton assemblages from eutrophic lakes

We conducted grazing experiments to test whether larger-bodiedDaphnia pulicaria have a different effect from smaller-bodiedDaphnia galeata mendotae on the composition of summer algalassemblages in eutrophic lakes. Three separate cubitainer experimentswere run for 5 days in a replicated factorial design utilizingtwo algal community types and the two Daphnia species. Inorganicphosphorus and nitrogen were added to prevent nutrient limitationof the algae. Both edible and inedible size fractions of chlorophylla increased in cubitainers without Daphnia spp. Grazer additionusually resulted in a reduction in edible chlorophyll; reductionswere greater in D.pulicaria cubitainers. Grazing by Daphniaspp. on presumed inedible chlorophyll was variable. Algal sizewas not always a good predictor of grazeability. The resultsof this study indicate that D.pulicaria, because of its greaterfiltration potential and ability to ingest larger particles,provides a stronger control on inedible-sized algae when comparedto equal numerical densities of D.g.mendotae. However, Aphanizomenonincreased as a response to heavy grazing pressure by D.pulicariaon other algal species. This suggests that biomanipulation effortsthat promote large-bodied Daphnia may not produce desirableresults if nutrient inputs remain high.     

Effects of Daphnia longispina on rotifer populations in a natural environment: relative importance of food limitation and interference competition

To test the relative importance of exploitative and interferencecompetition from Daphnia limiting rotifer populations in Bermejalesreservoir, birth rates of rotifers, food abundance, zooplanktonbiomass and size distribution of Daphnia were analysed fromhigh frequency sampling at two depths. A depletion in food abundanceby Daphnia feeding should be reflected in the birth rates ofrotifers susceptible to exploitative competition, while interferenceshould produce direct negative relationships between the numberof individuals of the interacting species. Results show thatsignificant relationships between food (chlorophyll-a) and birthrates of rotifers are found when the ratio chlorophyll-a/herbivorouszooplankton biomass is used as a measure of food availability.In our study, Anuraeopsis fissa was only affected through exploitativecompetition by Daphnia, while only Synchaeta oblonga showedevidence of being suppressed through interference competitionby this cladoceran.     

Influence of different sized Daphnia species on chlorophyll concentration and summer phytoplankton community structure in eutrophic Wisconsin lakes

We conducted field observations on 13 eutrophic Wisconsin lakesdominated by either the larger bodied Daphnia pulicaria or thesmaller bodied Daphnia galeata mendotae. While Daphnia numericaldensities were not significantly different between groups oflakes, pulicaria lakes had much higher Daphnia biomasses andfiltration potentials than galeata lakes. Although we foundsignificant differences in chlorophyll (Chl) a between bothgroups of lakes during June, on a seasonal basis populationsof different sized Daphnia were not associated with significantdifferences in Chl a. Filtration potential per se was the majordeterminant of Chl a, regardless of which Daphnia species dominated.However, in pulicaria lakes, the clear-water phase started earlier,lasted longer, and was usually characterized by greater Secchidisc readings than in galeata lakes. For large blue-green algaesuch as Aphanizomenon, D.pulicaria appeared to delay bloom conditions,but ultimately did not prevent the alga from growing. Our resultssuggest that high densities of large-sized Daphnia are a desirablegoal of biomanipulation because they can attain filtration potentialshigh enough to increase summer water clarity in eutrophic lakes.     

Clutch size and body size at first reproduction in Daphnia pulicaria at different levels of food and predation

Field data from seven alpine lakes in Serra da Estrela. Portugal.show that reproduction in Daphnia may be as efficiently controlledby fish predation and copepod predation on eggs in brood cavitiesas it is by food limitation. Body length and clutch size estimatesin Daphnia pulicaria revealed high inter- and intra-populationvariability in maturation size (body size at first reproduction).and in number of eggs per clutch. Daphnia at first maturationin lakes stocked with rainbow trout were half the size of thosefound in fishless lakes (body length of 0.86–0.95 and1.55–1.81 mm. respectively). The mean number of eggs perclutch was reduced to a similar degree by food limitation, predationby fish and copepod predation on eggs in brood cavities, butthe underlying mechanisms of this reduction were different.Food limitation caused smaller clutch sizes in all individuals,so variation remained the same. Fish predation caused the selectiveremoval of individuals with maximum clutches, so variation decreased.Copepod predation caused removal of eggs from brood cavitiesof randomly infested females, so that variation increased, particularlyat a high food level when non-infested females carried largeclutches of eggs.     

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.     

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.     

Finding the optimal vertical distribution: behavioural responses of Daphnia pulicaria to gradients of environmental factors and the presence of fish

1. The vertical distribution of zooplankton results from active habitat choice aiming to optimise fitness gain in a system of trade‐offs. 2. Using large, controlled indoor mesocosms (Plön Plankton Towers), we monitored the behavioural response of Daphnia pulicaria to vertical gradients of temperature, food, oxygen and light, in the presence and absence of fish predation. 3. In the absence of fish, Daphnia distributed as predicted by an ideal ‘free distribution with costs’. If the food was distributed homogeneously, they stayed in the warm epilimnion, while they balanced their time dwelling in epi‐ and hypolimnion if the food was concentrated in a deep‐water maximum. 4. However, oxygen depletion in the hypolimnion, representing an additional cost, prevented Daphnia from completely exploiting the hypolimnetic food maximum. Consequently, the proportion dwelling in the hypolimnion was larger if oxygen was not limiting. 5. Fish predation had an overwhelming effect, driving Daphnia into the hypolimnion under all experimental conditions. If permitted by oxygen availability, Daphnia used the whole hypolimnion, but oxygen depletion reduced their possible habitat to the upper hypolimnion with oxygen concentrations above c. 0.7 mg L^?1. As fish were less tolerant of low oxygen, the layer below the thermocline formed a predation refuge for Daphnia.     

Grazing and nutrient influences of Daphnia and Eudiaptomus on phytoplankton in laboratory microcosms

In two experiments, top-down and bottom-up influences of theherbivorous crustaceans Daphnia pulicaria and Eudiaptomus grac$$$loideson phytoplankton were compared in laboratory microcosms. Ina long-term experiment (63 days), both grazers were able toestablish populations. The Daphnia population exerted strongergrazing pressure, whereas Eudiaptomus fed more selectively.Daphnia retained relatively more phosphorus (P) and thus causedalgal P limitation; with Eudiaptomus as a grazer, both nitrogen(N) and P remained limiting. In a short-term experiment (1 day),N and P release rates and algal-specific grazing rates by bothconsumers were measured. In this experiment, the increase inconcentrations of soluble mineral nutrients (     

Chronic responses of different clones of Daphnia longispina (field and ephippia) to different food levels

Resting eggs are a fundamental reproductive strategy among freshwater cladocerans. Under adverse environmental conditions, whole Daphnia populations can disappear from a lake and a new community will arise from ephippial eggs. Since these new populations are subjected to genetic variation, their responses to environmental stress or contaminants can be different from the “original” population. In the present study, life history responses (reproduction and growth) of Daphnia longispina to different food concentrations was studied. Two Daphnia populations were tested: (a) field clones and (b) ephippial clones. Food (Selenastrum capricornutum) concentration was the stressor tested (absence of food, and low to high food concentrations). The results showed that reproductive responses of D. longispina to the tested food concentrations varied among clones, independently from their origin.