Temperature and the effects of elemental food quality on Daphnia

1. We examined the responses of two species of Daphnia to changes in food phosphorus (P) content, with animals reared at three different water temperatures. Specifically, we measured mass‐specific growth rate (MSGR), body P content and respiration rate of Daphnia magna and Daphnia pulex acclimatised to 10, 17.5 and 25 °C and fed food carbon : phosphorus (C : P) ratios of either 150 or 500. 2. The responses of these three physiological variables to temperature–food quality interactions were species‐specific. There was a significant interactive effect of temperature and food quality on D. magna, as the greatest proportional effect of food quality on growth was observed at 10 °C and reductions in body P because of low food P content were relatively greater at 25 °C. These effects may reflect the temperature dependence of mechanisms that reduce elemental constraints associated with food quality in D. magna. By contrast, there were no interactive effects between food quality and temperature on MSGR, body P or mass‐specific respiration of D. pulex. 3. It thus appears that temperature can alter food quality effects on Daphnia but the nature of these alterations depends upon the daphniid species and its thermal adaptability. Significant temperature–food quality interactions will complicate efforts to understand zooplankton nutrition in nature and warrant future consideration.     

Bacterial infection changes the elemental composition of Daphnia magna

1. An animal's elemental composition can be an important indicator of its physiological state and role in ecosystem nutrient cycling. We examined the interactive effects of bacterial (Pasteuria ramosa) infection and phosphorus (P)-poor food on the body stoichiometry of Daphnia magna. Daphnia were exposed to or held free of a bacterial parasite and fed algal food of different C:P ratios (100-500) over a 28-day period. 2. To assess the effects of exposure and infection on Daphnia stoichiometry, we measured their whole body content of carbon (C), nitrogen (N), and P on four different days (4, 8, 15, and 28) during the experiment. 3. We found strong effects of infection, food quality, and/or their interactions on the C, N, and P content of Daphnia, especially as the infectious disease progressed. At the end of the experiment, infected animals had significantly more C and less P in their bodies than uninfected conspecifics. Body N content of Daphnia consuming P-rich food was reduced by bacterial infection whereas Daphnia consuming P-poor algae showed increased body N content from infection. 4. Using a mass-balance model, we found that changes in N and P content of host bodies were largely accountable by disease-induced alterations to Daphnia reproduction (i.e. bacterial induced sterility) and the accumulation of Pasteuria spores in the body cavity. Our calculations also show that the observed increase in host C content could not be accounted for by loss of eggs or accumulation of bacterial spores. This instead must result from unidentified changes to underlying daphnid tissue C content. 5. These results demonstrate that intraspecific variation in zooplankton body stoichiometry can be caused by exposure to and infection by bacterial parasites. In addition, these effects were found to depend both upon the stage of the disease and upon the food quality consumed by the host.     

Protective effects of ectoine on heat-stressed Daphnia magna

Ectoine (ECT) is an amino acid produced and accumulated by halophilic bacteria in stressful conditions in order to prevent the loss of water from the cell. There is a lack of knowledge on the effects of ECT in heat-stressed aquatic animals. The purpose of our study was to determine the influence of ECT on Daphnia magna subjected to heat stress with two temperature gradients: 1 and 0.1 °C/min in the range of 23–42 °C. Time to immobilisation, survival during recovery, swimming performance, heart rate, thoracic limb movement and the levels of heat shock protein 70 kDa 1A (HSP70 1A), catalase (CAT) and nitric oxide species (NOx) were determined in ECT-exposed and unexposed daphnids; we showed protective effects of ECT on Daphnia magna subjected to heat stress. Time to immobilisation of daphnids exposed to ECT was longer when compared to the unexposed animals. Also, survival rate during the recovery of daphnids previously treated with ECT was higher. ECT significantly attenuated a rapid increase of mean swimming velocity which was elevated in the unexposed daphnids. Moreover, we observed elevation of thoracic limb movement and modulation of heart rate in ECT-exposed animals. HSP70 1A and CAT levels were reduced in the presence of ECT. On the other hand, NOx level was slightly elevated in both ECT-treated and unexposed daphnids, however slightly higher NOx level was found in ECT-treated animals. We conclude that the exposure to ectoine has thermoprotective effects on Daphnia magna, however their mechanisms are not associated with the induction of HSP70 1A.     

Effect of food concentration on the respiration of Daphnia magna

The relationship between respiration rate and dry weight forDaphnia magna was investigated in the absence of food. In addition the respiration and feeding rates of matureDaphnia magna females were measured simultaneously at differentChlorella concentrations. It was found that, besides the expected effect of food concentration on the feeding rate, there also was an effect of food concentration on the respiration rate. For food concentrations above a certain critical level a negative correlation between respiration rate and food concentration was found. Below this critical level the effect of the food concentration on the respiration rate was not significant.     

The effect of food on microparasite transmission in the waterflea Daphnia magna

Studies on dynamics of trophic transmission usually refer to the predation of animals that contain parasites. Here, in contrast, I study the transmission of external stages of parasites namely of spores of the microsporidian gut parasites Glugoides intestinalis and Micro1. These spores are filter fed by the waterflea Daphnia magna and I was particularly interested in the effect of food on transmission success. It was hypothesized that food has a double impact on transmission success because of increased spore uptake on the one hand and increased spore defecation of infected waterfleas on the other. I tested for these assumed food effects in exposure experiments where transmission could occur naturally from infected to uninfected waterfleas in the presence or absence of food.
The major assumptions were confirmed. In one system transmission success was increased fourfold in the presence of food and feeding infected hosts had lower spore loads. The other system showed evidence for these effects. Feeding activity correlated positively with food in both systems and spore loads in infected waterfleas partially determined spore loads in exposed waterfleas.
I discuss the impact of food on infected and uninfected waterfleas in particular and on transmission success and epidemiology in general.     

The interactive effects of temperature,food level and maternal phenotype on offspring size in Daphnia magna

Invertebrate offspring are usually larger in colder environments. To test for possible effects of covariates (e.g. maternal phenotype and feeding conditions) on this pattern, we performed a laboratory experiment to look at the effect of temperature on newborn weight in the planktonic crustacean Daphnia magna. Three tempèratures (12°C, 16°C and 22°C) and two food levels (10,000 cells ml^–1 and 150,000 cells ml^–1) were used, and offspring were examined from the first five clutches of mothers that had been maintained under the constant experimental conditions for three generations. Preliminary analysis suggested that newborn weight was significantly affected by temperature although patterns in the data were not clear cut. In addition, the covariates mother weight and clutch size were positively and negatively correlated with newborn weight, respectively; and later clutches tended to contain heavier offspring. Therefore, in an effort to control for the effects of the covariates, repeated-measures analysis of covariance was performed using ratio values of newborn weight/mother weight (relative newborn weight) as the dependent variable, clutch size as the covariate and clutch number as the repeated measures term. Now, temperature as a main effect in an ANCOVA model did not significantly influence relative newborn weight. The repeatedmeasure term clutch number also became nonsignificant, indicating that when differences in mother weight due to age were accounted for there were no overall differences in relative newborn weight between clutches from a particular mother. Temperature effects on relative newborn weight were only significant as part of interaction terms with food concentration and with clutch number. Thus there were different weight responses to temperature within food levels, and between clutch numbers within food levels. Under the low-food conditions newborn were heaviest at 16°C, lightest at 12°C and intermediate at 22°C. Conversely, under the high-food condition newborn were lightest at 16°C, heaviest at 12°C and again intermediate at 22°C. However, newborn tended to be heavier under the low food condition, and food concentration was highly significant as a main effect. Mother growth rate showed no significant relationship with newborn weight. It is concluded that direct temperature effects on relative newborn weight are marginal and nonsignificant. Temperature effects through interactions with food concentration and clutch number are important determinants of newborn weight, but relatively speaking account for only a small proportion of observed variance in newborn weight (25%), compared with the direct effect of food concentration (67%).     

Maternal food quantity affects offspring feeding rate in Daphnia magna

Maternal effects have wide-ranging effects on life-history traits. Here, using the crustacean Daphnia magna, we document a new effect: maternal food quantity affects offspring feeding rate, with low quantities of food triggering mothers to produce slow-feeding offspring. Such a change in the rate of resource acquisition has broad implications for population growth or dynamics and for interactions with, for instance, predators and parasites. This maternal effect can also explain the previously puzzling situation that the offspring of well-fed mothers, despite being smaller, grow and reproduce better than the offspring of food-starved mothers. As an additional source of variation in resource acquisition, this maternal effect may also influence relationships between life-history traits, i.e. trade-offs, and thus constraints on adaptation. Maternal nutrition has long-lasting effects on health and particularly diet-related traits in humans; finding an effect of maternal nutrition on offspring feeding rate in Daphnia highlights the utility of this organism as a powerful experimental model for exploring the relationship between maternal diet and offspring fitness.     

Maternal effects after sexual reproduction in Daphnia magna

In this study we investigated the relationship between maternalsize and size of ephippial hatchlings in Daphnia magna. We observedthat larger females produce larger ephippia; these larger ephippiahave a higher hatching probability, yield larger neonates, whichin turn yield larger adults, producing more eggs. We interpretthese observations as an explanation why older and larger femalesof D.magna continue to produce ephippia despite a higher predationrisk.     

Physiological effects of chronic silver exposure in Daphnia magna

Daphnia magna were exposed to a total concentration of 5.0+/-0.04 microg Ag/l, added as AgNO(3) (dissolved concentration, as defined by 0.45 microm filtration = 2.0+/-0.01 microg Ag/l) in moderately hard synthetic water under static conditions (total organic carbon = 4.80+/-1.32 mg/l) with daily feeding and water renewal, for 21 days. There was no mortality in control daphnids and 20% mortality in silver-exposed animals. Silver exposure caused a small but significant reduction of reproductive performance manifested as a 13.7% decrease in the number of neonates produced per adult per reproduction day over the 21-day exposure. However, silver exposed daphnids also exhibited a much more marked ionoregulatory disturbance, which was characterized by a 65% decrease in whole body Na(+) concentration, and an 81% inhibition of unidirectional whole body Na(+) uptake. Previous work on the acute toxicity of Ag(+) to daphnids has shown that Na(+) uptake inhibition is directly related to inhibition of Na(+),K(+)-ATPase activity. Therefore, we suggest that the Na(+) uptake inhibition seen in response to chronic silver exposure was explained by an inhibition of the Na(+) channels at the apical 'gill' membrane, since a 60% increase in whole body Na(+),K(+)-ATPase activity was observed in the chronically silver-exposed daphnids. Our findings demonstrate that, in broad view, the key mechanism involved in chronic silver toxicity in D. magna, the most acutely sensitive freshwater organism tested up to now, resembles that described for acute toxicity-i.e. ionoregulatory disturbance associated with inhibition of active Na(+) uptake, though the fine details may differ. Our results provide encouragement for future extension of the current acute version of the Biotic Ligand Model (BLM) to one that predicts chronic silver toxicity for environmental regulation and risk assessment. The results strongly suggest that Na(+) uptake inhibition is the best endpoint to determine sensitivity to both acute and chronic toxicity in the scope of future versions of the BLM for silver.     

Severe food stress has no detectable impact on developmental instability in Daphnia magna

Groups of individuals (three replicates of four treatment groups of 10 individuals) of Daphnia magna were exposed to nutritional stress in the forms of reduced food quantity and/or quality (phosphorus-limitation). Both types of stress led to reduced growth and reproduction, producing a gradient from surplus of high quality food to severe nutritional stress. There was no effect of nutritional stress on developmental instability measured either as fluctuating asymmetry or as phenotypic variability in body shape. Computer simulations confirmed that the lack of effect was not simply due to low statistical power. Low food quantity led to larger variability among individuals in overall body size, but this was most likely due to exploitative competition and competitive dominance of larger individuals. Low food quantity also led to increased occurrence of abnormal postabdominal spines (two spines appearing from one basis), but other analyses suggested that abnormal spines may actually represent an adaptive phenotypic response rather than a deviant phenotype caused by developmental instability. In conclusion, the results suggest that Daphnia magna are able to maintain high developmental stability even under extreme nutritional stress. This casts doubt on the utility of using any of the investigated measures of developmental instability as general indicators of environmental stress.     

Periphyton as alternative food source for the filter-feeding cladoceran Daphnia magna

1.  Daphnia magna , a well-studied primary consumer, is mainly known as a filter feeder. In this study, we investigated the ability of D. magna to use periphyton as an alternative food source to phytoplankton. We examined the development of laboratory populations fed with different food sources ( Desmodesmus subspicatus and/or periphyton or neither) over a period of 42 days, and observed the behaviour of the daphnids.
2.  The addition of periphyton to phytoplankton food led to an increase of daphnid population biomass. When fed with periphyton as the only food source, a small but stable D. magna population developed.
3.  The behaviour of daphnids fed with both food sources revealed a preference for feeding on D. subspicatus . Only below a concentration of D. subspicatus of approximately 0.05 mg C L⁻¹ (0.4 × 10⁷ cells L⁻¹) did D. magna use periphyton as an alternative food source.
4.  Periphyton showed distinct reactions to grazing by D. magna . The thickness of the periphyton layer was reduced from about 4 to 1 mm and we observed a change in species composition due to grazing.
5.  The ability of D. magna to graze on periphyton could serve to stabilize its population density and reinforce its competitive advantage over other cladocerans. By switching between food sources, D. magna can act as a coupler between pelagic and benthic habitats and food webs.     

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.     

Long-term temperature acclimation in Daphnia magna: effects on filtering rates

Populations of a laboratoiy clone of Daphnia magna were acclimatedat 5, 10, 15 and 24C, and a varying temperature regin for aminimum of 70 days. The effect of temperature (5–25C)on filtering rates was measured for animals acclimated to theconstant temperatures and for a wild population collected froma reservoir. Acclimation temperature strongly influenced ffltcrmgrates at test temperatures. Animals with acclimation temperaturesclosest to the test temperature tended to have highest filteringrates at that temperature.     

Temperature reaction norms of Daphnia magna: the effect of food concentration

1. The effects of temperature and food concentration on the fitness of Daphnia magna were tested in a 4×4 factorial flow-through design. Food ranged from 0.1 to 1.0 mg C L⁻¹ and temperature ranged from 15 to 30 °C. 2. The juvenile growth rate ( g _j) was used to construct reaction norms for temperature at varying food concentrations. Two clones isolated from the same pond at different seasons did not differ with respect to their temperature responses. Reaction norms had the shape of an optimum curve with highest values around 20 °C. There was a significant temperature–food interaction as the temperature response was most pronounced when the food was not limiting. 3. Differences in fitness were a consequence of different responses of physiological parameters to food and temperature. Age and size at first reproduction, as well as egg numbers, decreased with increasing temperature and decreasing food concentration. 4. As the temperature effect was strongest at the highest food concentrations, it can be concluded that environmental warming may affect D. magna more through a temperature rise earlier in spring rather than in summer.     

Response of the respiratory rate of Daphnia magna to changing food conditions

Summary The respiratory rate of the cladoceran Daphnia magna was measured at varying concentrations of a green alga in a flow-through respirometer. Daphnids were either preconditioned to the respective food concentration or the food concentration was suddenly changed during the experiment and the response of the respiratory rate monitored. Previously starved animals were provided with food or prefed animals were deprived of food. The respiratory rate increased considerably with increasing concentrations of algae until a maximum rate was reached at a critical algal concentration corresponding to the incipient limiting level for feeding. The response of the respiratory rate to changing food conditions was fast, lagging only a few minutes behind the change in food. The respiratory rate of starved daphnids increased quickly when they received food, even at low concentrations. Prefed daphnids responded to the replacement of the food suspension by filtered water with reductions in their respiratory rates. A linear relationship between the assimilation rate and the respiration rate was found, indicating that the more than twofold range of the respiratory rate was due to specific dynamic action. The SDA coefficients of 0.15–0.24 found for Daphnia are consistent with values for marine zooplankton.     

Effect of food concentration on the chronic toxicity of sodium dodecyl sulphate to Daphnia magna

Food concentration supplied during subchronic and chronic toxicity tests, is one of the most important factors that might influence the response of test organisms to toxicants. The green microalga Scenedesmus incrassatulus was used as food for the cladoceran Daphnia magna, in a chronic toxicity test with the toxicant sodium dodecyl sulphate (SDS). Test concentrations were 0.625, 1.25, and 2.5 mg l^-1 of SDS, equivalent to 1/40, 1/20, and 1/10 of the average 48-h LC50 previously determined. Food concentrations were 9.5, 19, and 38 mg l^-1 (dry wt.). Survival and reproduction were recorded, and the data were analyzed using a life-table approach. After 55 days, the main findings were as follows:

Effect of food type on the acute and chronic toxicity of copper to Daphnia magna

SUMMARY. Daphnia magna was maintained on two diets and exposed to acute and chronic copper stresses. Animals fed vitamin-enriched algae were less sensitive to a chronic copper stress than animals fed a trout-granule diet. This conclusion is based on the responses of cohorts as measured by survival, mean brood size and r , the instantaneous rate of population growth. Application factors were also significantly different for animals reared on the two foods (0.47 for algal-fed animals, 0.12 for animals reared on trout-granules). In contrast to chronic toxicity, 72 h LC₅₀ values were not significantly different for animals maintained on the two foods. This suggests that the mechanism of toxic action is different for acute and chronic toxicity and that, if so, the ratio between chronic and acute toxicity would not be a constant under different environmental conditions.     

The heart of Daphnia magna: effects of four cardioactive drugs

We used Daphnia magna bioassays to determine the LC(50) and the effects on the heart of the cardioactive drugs ouabain, verapamil, metaproterenol and metoprolol. Distinctions were made between the pharmacological and toxicological effects of these drugs and the adequacy of physicochemical characteristics of its habitat (reconstituted water). Video microscopy and digital image processing were used to study the pharmacological effects on the heart. D. magna exhibited the expected sensitivity to the reference toxicant sodium dodecyl sulfate with a LC(50) of 15.6+/-4.5 mg/l. All drugs were toxic with 48 h-LC(50) of 2.03 mg/l ouabain, 7.04 mg/l verapamil, 32.45 mg/l metaproterenol and 76.21 mg/l metoprolol. Ouabain was the most toxic and caused a positive concentration-dependent inotropic effect. Verapamil caused positive chronotropic and inotropic effects, while metaproterenol showed positive concentration-dependent chronotropic effects at high concentrations (10(-3) and 10(-4) M). Metoprolol induced a positive chronotropic effect at low concentrations (10(-8), 10(-7), 10(-6) M) and a negative chronotropic effect at high concentration (10(-4) M). Ouabain, metaproterenol and metoprolol in D. magna caused similar effects to those produced in mammals. In contrast, verapamil caused opposite effects. The results suggest the presence of Na(+), K(+)-ATPase receptors to verapamil and of non-specific adrenergic receptors in heart of D. magna.