Morphological adaptations in filtering screens of Daphnia galeata to food quantity and food quality

We reared clones of the waterflea Daphnia galeata, a common grazer in many types of lakes, under several food regimes to study adaptations to feeding conditions in filter screen morphology and life history. As food regimes, we used low and high concentrations of the green alga Scenedesmus, a high concentration of the filamentous cyanobacterium Oscillatoria, and a mixture of Scenedesmus and clay. Natural seston from Lake IJsselmeer was also tested. Clones from two contrasting habitats (mesotrophic versus hypertrophic lake) did not differ in either population growth rate, r, or filtering structures. However, all the clones exhibited phenotypic plasticity when reared under low and high food concentrations. In relation to their body length, daphnids grew larger filter screens with longer setae in response to low food concentrations, thus enhancing filtering efficiency. Supplementing the low food concentration with inert clay particles had no effect on either the growth performance or filter screen morphology. In both cases, the growth and reproduction were low and filtering areas large. Filter screens were of intermediate size when daphnids were supplied with food supporting intermediate growth and reproduction. From these laboratory results, we concluded that the nutritional status of Daphnia is a more important cue for the phenotypic response of the filter screen morphology than particle concentration.      

Seasonal patterns of food limitation in Daphnia galeata: separating food quantity and food quality effects

Food limitation for the cladoceran Daphnia galeta was measuredgrowing them with natural lake seston for one growing seasonunder standardized conditions in the laboratory. Growth rateswere related to several measures of food quantity. Particulateorganic carbon (POC) <30 µm was the best predictorof total food availability measured. It was better than chlorophyll-a(Chl-a) of the same size fraction, algal volume calculated fromcell counts or total particle volume from Coulter Counter measurements.Daphnids were also grown with Scenedesmus acutus under the sameconditions and their growth rates were compared. Sestonic carbonconcentrations were related to Scenedesmus carbon concentrations,that provided equal growth rates. The ratio of the carbon concentrationsof both food types (seston, Scenedesmus) represented the foodquality of seston. Thus, the nutritional quality of naturallake seston relative to a standard alga was determined overone growing season. Growth rates of seston fed animals revealedthat they were food limited during long periods of the year.This was also ascertained by standard clutch sizes of fieldanimals. However, the causes of food limitation varied withinthe duration of the studied period. Whereas during the clear-waterphase the food quantity was limiting (while the quality washigh: 100% Scenedesmus), shortly after on 10 June and 17 Junefood conditions were close to threshold concentrations due tolow food quality. For the remainder of the summer, a POC of0.3–0.6 mg C I^–1 with a quality of     

Phytoplankton responses to grazing by Daphnia galeata in the biomanipulated Bautzen reservoir

We studied the response of phytoplankton to grazing by Daphnia galeata in the hypertrophic Bautzen reservoir (Dresden, Germany) from January 1995 to May 1996 during a long-term whole-lake biomanipulation experiment. The correlation between clearance rate of D. galeata and total phytoplankton biomass was negative only if biomass of Microcystis aeruginosa was excluded. This suggests that M. aeruginosa is the main grazing resistant phytoplankton species in the Bautzen reservoir. Except for M. aeruginosa and grazing-resistant Staurastrum quadridentatum spec. nov. (Scharf, 1995) no other phytoplankton species was able to reach a biovolume above 1 mm3 L-1 when the clearance rate of Daphnia exceeded 0.1 L L-1 d-1. There was significant positive correlation between mean cell or colony size of phytoplankters and clearance rate of D. galeata, showing an advantage of bigger cells or colonies at high grazing pressure. Cross-correlation indicated a time lag of about one month between changes in grazing pressure and a change in phytoplankton mean size. The phytoplankton species were divided into edible and inedible fractions, depending on their width and length. No edible species were able to reach high biovolumes during high biomasses of D. galeata but a positive correlation was found between the edible fraction of phytoplankton biovolume and the clearance rate of D. galeata. However, this relationship disappeared when the ‘ingestible edible’ fraction of M. aeruginosa was excluded, suggesting a rejection of ‘edible’ Microcystis colonies by D. galeata. A negative correlation was found between the inedible fraction of phytoplankton biovolume without M. aeruginosa and the clearance rate of D. galeata which might be due to superior competition of M. aeruginosa. We could clearly show that biomanipulation might not work well with respect to the reduction of total phytoplankton biomass under hypertrophic conditions and finally discuss a theoretical threshold of phosphorus (probably around 80 μg L-1), below which biomanipulation should become effective. This revised version was published online in July 2006 with corrections to the Cover Date.     

Effect of food conditions on the bacterial feeding of Daphnia galeata

Feeding selectivities of Daphnia galeata were measured using various density combinations of ¹⁴C-labeled Chlamydomonas and ³H-labeled natural bacteria. In all food combinations, D. galeata preferred algae. Selectivity coefficient (bacterial clearance rate: algal clearance rate) was 0.35 at the level of incipient food limitation (1.5 × 10⁶ µm³ ml^–1).However, it increased up to 0.9 with increasing total food concentration. A simple mechanical sieving model alone cannot explain such a change in the selectivity coefficient. A mechanism of clogging suggestively explain the increase of bacterial ingestion efficiency by D. galeata, especially at high food concentrations. The change of selectivity coefficient related to food condition suggests that bacterial feeding by the Daphnia is incidental event associated with the algal feeding.     

Synergistic effects of different food species on life-history traits of Daphnia galeata

In this paper we describe the life history consequences of feeding Daphnia galeata with different food types in different concentrations. We fed the animals with four concentrations of two green algae Scenedesmus obliquus and Chlamydomonas globosa, given separately as well as in a 1:1 mixture. Growth, reproduction and condition of the animals and consequently the intrinsic rate of population increase, r, increased with the concentration of the food for all three experimental series. The life history parameters measured were lowest in the animals raised on C. globosa. Animals fed on S. obliquus showed higher values for r. We observed a distinct synergistic effect of the two algal species: growth, reproduction, condition and r were significantly higher for the animals raised on the mixed media as compared to the animals raised on the mono-algal media. We conclude that the two algal species represented complementary food resources.     

Reduction of the filtration rate of Daphnia galeata by dissolved photosynthetic products of edible phytoplankton

The filtration rate of Daphnia galeata was determined in in situ experiments in Bautzen Reservoir and in laboratory experiments, where daphnids were exposed to filtrates that previously contained either natural phytoplankton or cultured eukaryotic algae (Scenedesmus obliquus or Asterionella formosa), respectively. Individual filtration rate (FR) was measured using fluorescent beads, taking into account ingested beads in the gut only. Compared to heated control treatments (100 °C), dissolved compounds released by the nutritious cultured algae during the preconditioning phase or by the natural phytoplankton assemblages from Bautzen Reservoir strongly reduced the filtration rate of D. galeata (down to 60%). Heating deactivated these dissolved compounds. A significant correlation was found between primary production measured in situ and the reduction of FR in the filtrate of reservoir water, indicating that extra-cellular products released during photosynthesis triggered the reduction of the filtration rate. The ratio of ingested to collected beads was used to quantify the proportion of food, which was not only collected but passed the mouth of D. galeata. The ratio of ingestion to collection was compared between filtered and unfiltered reservoir water both media identical with respect to the concentration of dissolved compounds, whereas other factors (e.g. food concentration, temperature, filtration rate) were different. The changes in this ratio between filtered and unfiltered reservoir water suggest that D. galeata is capable of a chemosensory control of the ingestion behaviour by detecting external metabolites.     

Tolerance to low ambient calciumshows inter-population differences in Daphnia galeata

The tolerance to low ambient calcium (Ca) was assessed in twopopulations of Daphnia galeata. There was a threshold for survivalbetween 0 and 2 mg Ca l^-1. However, the ability to cope withlow Ca concentrations clearly differed between the two populations,as the population from the low-Ca locality was less tolerantto low Ca. Additionally, neonate individuals had poorer survivalthan adults when reared at ~0 mg Ca l^-1, supporting a suggestedjuvenile bottleneck regarding the tolerance to low Ca concentrations.The mean specific Ca content in adult D. galeata was not differentbetween the two populations, but individuals reared in mediumwith 1 mg Ca l^-1 only had two-thirds of the Ca content of thosereared in medium with 10 mg Ca l^-1. The significant differencesbetween the two populations investigated suggest that inter-populationvariation in tolerance to low ambient Ca concentrations mightbe important to explain the success of Ca-demanding crustaceansin softwater lakes.     

An in situ test of the effects of food quality on Daphnia population growth

The temporal variation of the surface deposit feeders in a mudflat of Chuwei Mangrove Forest showed a significant input of populations from the recruitments in the spring. The chemical analysis of the sediment in the top 8 cm showed that the sedimentary organic content had a subsurface maximum value in the deeper layer in most cases. Most of the temporal variations in the organic carbon and nitrogen contents occurred in the top few centimeters. The high C:N ratio and low organic nitrogen content in the sediment suggests the supply of carbon (or energy) to the benthos was low. The increase of labile protein content in the deep layer might be due to the downcore transport of sediment nutrients by the bioturbation. Laboratory growth experiment suggests that some the chemical characters examined in this study may represent the sediment nutrients to the animals.     

Genetic Structure of Daphnia galeata Populations in Eastern China

This study presents the first examination of the genetic structure of Daphnia longispina complex populations in Eastern China. Only one species, D. galeata, was present across the eight investigated lakes; as identified by taxon assignment using allelic variation at 15 microsatellite loci. Three genetically differentiated D. galeata subgroups emerged independent of the type of statistical analysis applied. Thus, Bayesian clustering, discriminant analysis based on results from factorial correspondence analysis, and UPGMA clustering consistently showed that populations from two neighbouring lakes were genetically separated from a mixture of genotypes found in other lakes, which formed another two subgroups. Clonal diversity was high in all D. galeata populations, and most samples showed no deviation from Hardy-Weinberg equilibrium, indicating that clonal selection had little effect on the genetic diversity. Overall, populations did not cluster by geographical origin. Further studies will show if the observed pattern can be explained by natural colonization processes or by recent anthropogenic impact on predominantly artificial lakes.     

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     

Reproductive potential of Daphnia galeata in relation to food conditions: implications of a changing size-structure of the population

A field study was conducted in the highly eutrophic, biomanipulated Bautzen Reservoir to investigate the influences of food conditions on fecundity of Daphnia galeata during spring and early summer in four consecutive years. By defining a relation between body size and maximum brood size, the realised reproductive potential (RRP) of Daphnia was expressed as percentage of actual egg numbers of the population in relation to calculated maximum egg numbers. This parameter is independent of the size structure of the population. It was found to correlate generally with mean clutch size. However, deviations occurred if size at maturity (SAM) declined. Extremely low SAM values (0.8 mm) were recorded in 2 years, corresponding with increasing food concentrations. Low clutch sizes in these situations can be explained by small body sizes of ovigerous females, whereas RRP increased, thus reflecting good food conditions. From the comparison of different reproductive parameters, it is concluded that RRP is the most reliable one with regard to bottom-up influences in case of extremely variable size structure and possible predation impacts. Fecundity of D. galeata was clearly food limited during the clear-water stage. Relatively high POC-threshold concentrations of 0.26 mg C l^–1 suggest that food quality was low and may be equally important as food quantity in limiting egg production.     

Application of individual growth and population models of Daphnia pulex to Daphnia magna, Daphnia galeata and Bosmina longirostris

Daphnia models for individual growth and population dynamics have been developed in the manner of models developed by Gurney, McCauley, Andersen and others. All or most of the earlier models were parameterized for Daphnia pulex; we have used the D. pulex model as a baseline model for other species of Daphnia such as magna, galeata and also Bosmina longirostris. Because of the lack of ample data for D. magna, D. galeata and B. longirostris, some of the physiological data had to be relied on the other species whose data were available and in some case calibrated. We were able to produce reasonable results for individual growth as well as population dynamics under the controlled laboratory conditions. Most of the results were compared with the available laboratory data for population as well as growth. All the simulations have been done under high and low food concentrations. The animals are assumed to be feeding on green algae (Chlamydomonas reinhardtti) under the laboratory conditions of 18–20°C. The continuous growth until the end of the life was observed in smaller B. longirostris, whereas rapid growth in the beginning and slower after the start of the reproduction was observed in Daphnia species. The smaller species matured earlier than larger species. B. longirostris population sustained better than Daphnia species in medium food concentrations.     

Effects of neonate size and food concentration on the life history responses of a clone of the hybrid Daphnia hyalina × galeata to fish kairomones

1. The influence of fish kairomones on the life history parameters of a clone of the hybrid Daphnia galeata × hyalina was measured in laboratory experiments with small and large neonates at one food concentration. The influence of fish kairomones on neonates of the same size at two food concentrations was also investigated.
2. Neonate size (maternal investment per egg) changed the influence of fish kairomones on life history parameters. Daphnids that were small at birth showed greater differences between fish (kairomones present) and no-fish (kairomones absent) treatments for age at maturity and number of eggs in the first clutch than animals that were large at birth. On the other hand, animals that were large at birth differed more in size at maturity between fish and no-fish treatments than small neonates.
3. The food concentrations used in the experiments (0.25–0.75 mg C l^–1) hardly affected the influence of fish kairomones on the life history characteristics studied.
4. The effect of fish kairomones on the size of neonates from the first clutch was similar for large and small daphnids. In the no-fish treatment significant differences were found between small and large neonates for age at maturity and number of eggs in the first clutch. In the fish treatment size at maturity differed significantly between small and large neonates.
5. In life history experiments, neonate size has to be taken into account when measuring the effects of fish kairomones because size at birth influences the results.     

Swimming of Daphnia galeata x hyalina in response to changing light intensities: Influence of food availability and predator kairomone

Experiments showed that phototactic downward swimming in Daphnia galeata x hyalina as caused by a relative increase in light intensity (stimulus) is influenced by predator kairomone and food availability. The swimming responses at four different combinations of food availability and fish kairomone were analysed. Addition of both food and kairomone led to a significant increase in percentage of animals that responded to the light stimulus, but there was no significant interaction effect.We also found that kairomone and food had significant impact on displacement velocity and on the time between start of the stimulus and onset of the response.     

Effects of neonate size and food concentration on the life history responses of a clone of the hybrid Daphnia hyalina × galeata to fish kairomones

1. The influence of fish kairomones on the life history parameters of a clone of the hybrid Daphnia galeata × hyalina was measured in laboratory experiments with small and large neonates at one food concentration. The influence of fish kairomones on neonates of the same size at two food concentrations was also investigated.
2. Neonate size (maternal investment per egg) changed the influence of fish kairomones on life history parameters. Daphnids that were small at birth showed greater differences between fish (kairomones present) and no-fish (kairomones absent) treatments for age at maturity and number of eggs in the first clutch than animals that were large at birth. On the other hand, animals that were large at birth differed more in size at maturity between fish and no-fish treatments than small neonates.
3. The food concentrations used in the experiments (0.25–0.75 mg C l^–1) hardly affected the influence of fish kairomones on the life history characteristics studied.
4. The effect of fish kairomones on the size of neonates from the first clutch was similar for large and small daphnids. In the no-fish treatment significant differences were found between small and large neonates for age at maturity and number of eggs in the first clutch. In the fish treatment size at maturity differed significantly between small and large neonates.
5. In life history experiments, neonate size has to be taken into account when measuring the effects of fish kairomones because size at birth influences the results.     

Influence of food density on respiration rate of two crustacean plankters,Daphnia galeata and Bosmina longirostris

Summary The influence of food density on respiration rate was measured for two cladoceran plankton species, Daphnia galeata and Bosmina longirostris, over the range 0 to 2.5 mg C 1^-1, using the modified Winkler technique in order to examine how this affects the respiration rate and whether the functional response is the same in the two species. The respiration rate for animals of equivalent body size did not differ significantly between the two species in the absence of food, but was significantly lower in Bosmina longirostris than Daphnia galeata at high food density. Within a species, the response of respiration rate to changing food density did not differ among individuals of different body size. The respiration rate of D. galeata increased with increasing food density and reached a plateau at a high food density. A similar response curve was also found with the respiration rate of B. longirostris, although the response was less conspicuous. This response curve cannot be explained by the energy cost of known feeding behavior in cladocerans. Since the respiration rate related linearly with the assimilation rate, increase in food density seemed to increase the respiration rate by increasing the energetic cost required to process food biochemically, known as specific dynamic action.     

Feeding behaviour and morphology of filtering combs of Daphnia galeata

The population of Daphnia galeata Sars from the fish pond Velký Pálenec (Blatná, Czechoslovakia) living in high food conditions (7 mgC l^–1) was characterized by a small size of the filtering comb on the thoracic limb 3, measured as seta length, length of the base of the comb and number of setae (population 1). One month cultivation of this population in low food conditions (1.5 mgC l^–1) in the laboratory resulted in twofold increase in size of the filtering comb (population 2). Filtering and ingestion rates of both populations were measured at eight concentrations of food (approximately 0.025–3.2 mgC l^–1) using ¹⁴C labeled Scenedesmus acutus. The results show that size of the filtering combs influences considerably feeding behavior of Daphnia. The comparison of animals with the same body length suggests that the population with a large comb feeds at concentration of food below 0.4 mgC l^–1 more intensively and reaches the maximum of the filtering rate at a lower concentration than the population with a small comb. The situation is opposite at concentration above 0.4 mgC l^–1. The higher values of theoretical flow in population with a small projection of filtering area suggest that this population has to compensate disadvantage of a small comb with the higher appendages beat frequency.     

Quaternary diversification in a sexual Holarctic zooplankter, Daphnia galeata

The effects of Quaternary glacial range partitioning on the diversification of Holarctic biota remain unclear. Glacial refugial lineages may form vicariant species, hybrid products, or merge after secondary contact. Here, we assess the effects of Quaternary glaciation on a Holarctic sexual zooplankter, Daphnia galeata, with apparently marked dispersal capacity and a widespread hybrid lineage in the New World. We collected samples of this species from 148 Holarctic lakes, analysed the nuclear and mitochondrial gene sequences, and tested predictions for hypotheses that account for the origin and spread of the New World D. galeata. We detected five nuclear phylogroups and four mitochondrial phylogroups, most of which were restricted to either the New World or the Old World. The oldest mitochondrial phylogroup was restricted to Japan. One major mitochondrial clade was distributed throughout the Holarctic, but only four haplotypes were shared among continents, and analysis of molecular variance indicated significant structure at the continental level. Haplotype sharing among continents could largely be attributed to anthropogenic introductions. Mismatch distributions, haplotype networks, phylogenetic trees, longitudinal haplotype diversity erosion and coalescence analyses are consistent with colonization from an Old World and a New World refugium. Our nuclear and mitochondrial DNA sequence evidence supports the hypothesis that the New World D. galeata underwent introgression with Daphnia dentifera, with dispersal being enhanced by glaciation. We conclude that Quaternary glaciation had a pronounced effect on the diversification of a Holarctic sexual zooplankter.     

Influence of food quality on growth and reproduction in Daphnia

SUMMARY. 1. Growth and reproduction were measured for Daphnia pulex (Leydig) fed Scenedesmus obliquus (Turp.) Kutz. grown with nitrogen-sufficient and nitrogen-deficient media. The incorporation of carbon into the body of D. pulex was traced using ¹⁴C-labelled algae and biochemical fractionation.
2. Scenedesmus cultured on N-defkient media allocated more carbon to lipid and less to protein than when cultured on N-sufficient media.
3. In a short-term feeding experiment, Daphnia accumulated more lipid when fed N-deficient algal cells high in lipid content than when fed algae grown on N-sufficient media that were lower in lipid.
4. Animals grew faster and produced larger broods when fed algae grown on an N-sufficient medium over an 8-day period. Daphnids fed N-deficient algae had higher quantities of body lipid, suggesting it was not a lack of energy which limited their growth and reproduction.
5. Daphnia fed algae grown on N-deficient media allocated more lipid to eggs, and their offspring lived longer under starvation conditions.