Divergent phenotypic responses to predators and cyanobacteria in Daphnia lumholtzi

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Habitat partitioning of native and exotic Daphnia in gradients of temperature and food: mesocosm experiments

1. Invasion of tropical zooplankton into temperate lakes provides an interesting opportunity to explore habitat segregation in a thermal gradient. 2. We explored differing vertical positioning of native and exotic Daphnia (Daphnia mendotae and Daphnia lumholtzi) in a large indoor mesocosm system (Plön plankton towers) during 2 month‐long experiments. The two towers were manipulated to provide gradients of both temperature (15–29 °C) and algal food (0.05–0.58 mg C L^?1) and a day–night cycle. 3. Both juvenile and adult D. lumholtzi showed a ‘typical’ vertical migration pattern, with higher densities in the epilimnion at night than during the day. They avoided the food‐poor middle layer. In contrast, D. mendotae adults showed little tendency to migrate into the epilimnion at night, remaining in the cooler hypolimnion while juveniles migrated. The vertical distribution of D. mendotae appeared unaffected by the presence of D. lumholtzi. 4. The strong migration behaviour of D. lumholtzi in the absence of fish cues suggests that this behaviour may be a constitutive trait. Habitat partitioning of the two species is probably the result of different thermal tolerances, with D. mendotae constrained to remaining in deeper water by high temperatures in the epilimnion and the tropical D. lumholtzi able to use the warm epilimnion at night.     

Linking biotic interactions and climate change to the success of exotic Daphnia lumholtzi

1. Warmer temperatures may increase cyanobacterial blooms in freshwater ecosystems, yet few ecological studies examine how increases in temperature and cyanobacterial blooms will alter the performance of non‐native species. We evaluated how competitive interactions and interactions between these two drivers of ecological change influence the performance of non‐native species using the native zooplankton Daphnia pulex and the non‐native zooplankton Daphnia lumholtzi as a model system. Based on the literature, we hypothesised that D. lumholtzi would perform best in warmer temperatures and in the presence of cyanobacteria. 2. Laboratory competition experiments showed that in the absence of competitors, growth rates of D. pulex (but not D. lumholtzi) were reduced at higher temperatures and with the cyanobacterial foods Anabaena flos‐aquae and Microcystis aeruginosa. In the presence of competitors, however, D. pulex emerged as the superior resource competitor at both temperatures with cyanobacterial food. We therefore predicted that, if competitive interactions are important to its establishment, D. lumholtzi would perform best in the absence of cyanobacteria in heated environments. 3. As predicted, when both species were introduced at low densities in field mesocosms, D. lumholtzi performed best at high temperatures without added cyanobacteria and worst at ambient temperatures with added cyanobacteria, indicating that competitive interactions are likely to be important for its establishment. 4. Taken together, these studies indicate that, while D. lumholtzi may benefit from increases in temperature, associated increased cyanobacterial blooms may hinder its performance. Thus, our findings underscore the importance of considering biotic interactions such as competition when predicting the future establishment of non‐native species in response to climate warming.     

The effects of Daphnia on nutrient stoichiometry and filamentous cyanobacteria: a mesocosm experiment in a eutrophic lake

1. Stoichiometric theory predicts that the nitrogen : phosphorus (N : P) ratio of recycled nutrients should increase when P‐rich zooplankton such as Daphnia become dominant. We used an enclosure study to test the hypothesis that an increased biomass of Daphnia will increase the relative availability of N versus P sufficiently to decrease the abundance of filamentous cyanobacteria. The experiment was conducted in artificially enriched Lake 227 (L227) in the Experimental Lakes Area (ELA), north‐western Ontario, Canada. Previous studies in L227 have shown that the dominance of filamentous, N‐fixing cyanobacteria is strongly affected by changes in the relative loading rates of N and P. 2. We used a 2 × 2 factorial design with the addition or absence of D. pulicaria and high or low relative loading rates of N and P (+NH₄, –NH₄) in small enclosures as treatment variables. If Daphnia can strongly affect filamentous cyanobacteria by altering N and P availability, these impacts should be greatest with low external N : P loading rates. The phytoplankton community of L227 was predominantly composed of filamentous Aphanizomenon spp. at the start of the experiment. 3. Daphnia strongly reduced filamentous cyanobacterial density in all enclosures to which they were added. The addition of NH₄ had only a small impact on algal community composition. Hence, we conclude that Daphnia did not cause reductions in cyanobacteria by altering the N : P ratio of available nutrients. 4. Despite the lack of evidence that Daphnia affected filamentous cyanobacteria by altering the relative availability of N and P, we found changes in nutrient cycling consistent with other aspects of stoichiometric theory. In the presence of Daphnia, total P in the water column decreased because of an increase in P sedimentation. In contrast to P, a decrease in suspended particulate N was offset by an increase in dissolved N (especially NH₄). Hence, dissolved and total N : P ratios in the water column increased with Daphnia as a result of differences in the fate of suspended particulate N versus P. There was minimal accumulation and storage of P in Daphnia biomass in the enclosures. 5. Our experiment demonstrated that Daphnia can strongly limit filamentous cyanobacterial abundance and affect the biogeochemical cycling of nutrients. In our study, changes in nutrient cycling were apparently insufficient to cause the changes in phytoplankton community composition that we observed. Daphnia therefore limited filamentous cyanobacteria by other mechanisms.     

Evolution of life histories and yields in experimental populations of Daphnia magna

An experiment on life-history evolution is described in which replicated populations of the Cladoceran Daphnia magna , made up of a standard mixture of clones, were subjected to two contrasting culling regimes, involving removal of small or large individuals. After approximately 150 days of culling, analysis of the life histories showed that genetic differences had emerged between culling regimes. Clones selected by culling small sized individuals grew rapidly through small size classes, whereas those selected by culling large sizes grew slowly through small size classes, with the result that the age at which they became vulnerable to harvesting was delayed. In addition, there was some redistribution of reproduction towards size classes that were not culled. This evolution is consistent with a major decline observed in the yield from populations in which large individuals were culled. We argue that changes of this general kind are likely to take place in exploited populations and that serious consideration should be given to evolutionary aspects of the management of such populations.     

Negative effects of UVB-irradiated phytoplankton on life history traits and fitness of Daphnia magna

SUMMARY 1. We tested the effect of ultraviolet-B (UVB)-irradiated phytoplankton on life history characteristics of Daphnia magna . Two phytoplankton species were used, Chlamydomonas reinhardtii and Cryptomonas pyrenoidifera . The phytoplankton species were cultured under photosynthetically active radiation (PAR) conditions, and under PAR supplemented with ultraviolet-A and ultraviolet-B radiation, and fed to Daphnia .
2. Life history traits of Daphnia were negatively affected when fed on UVB-irradiated Cryptomonas . Size at maturity was depressed and fewer juveniles with lower fitness were produced in the UVB treatments. In the Chlamydomonas experiment, no significant effects were found.
3. The cause of the observed UVB effects is likely to be constraints in food quality. Ultraviolet-B radiation thus has the potential of inhibiting energy transfer from the first to the second trophic level.     

Measuring Daphnia life history in the wild: The efficacy of individual field cages

Life‐history studies are often conducted in a laboratory environment where it is easy to assay individual animals. However, factors such as temperature, photoperiod, and nutrition vary greatly between laboratory and field environments, making it difficult to compare results. Consequently, there is a need to study individual life histories in the field, but this is currently difficult in systems such as Daphnia where it is not possible to mark and track individual animals. Here, we present a proof of principle study showing that field cages are a reliable method for collecting individual‐level life‐history data in Daphnia magna. As a first step, we compared the life history of paired animals reared outside and inside cages to test the hypothesis that cages allow free flow of algal food resources. We then used a seminatural mesocosm setting to compare the performance of individual field cages versus glass jars refilled with mesocosm water each day. We found that cages did not inhibit food flow and that differences in life histories between three clones detected in the jar assays were also detectable using the much less labor‐intensive field cages. We conclude that field cages are a feasible approach for collecting individual‐level life‐history data in systems such as Daphnia where individual animals cannot be marked and tracked.     

Life history of Daphnia galeata in a hypertrophic reservoir and consequences of non-consumptive mortality for the initiation of a midsummer decline

SUMMARY 1. Field and laboratory investigations were combined in a 2‐year study on the initiation of a midsummer decline of Daphnia galeata Sars in a hypertrophic reservoir. Quantitative field samples were taken twice a week, and, adult and juvenile mortality rates were calculated. Patterns of reproduction and survival of daphnids born during spring and early summer under fluctuating food conditions were determined in life‐table experiments. 2. The abundance of Daphnia increased strongly in early May and declined in June 1998 (midsummer decline). In 1999, Daphnia density increased only slowly in spring and remained constantly high throughout the summer. 3. Food conditions (concentrations of POC_{<30 μm}) for daphnids deteriorated in both years in response to increasing Daphnia densities, resulting in a clear‐water phase of about 4 weeks. When Daphnia abundance declined in 1998, POC_{<30 μm} concentrations increased greatly, whereas in 1999 food conditions improved only slightly and Secchi depth remained high. 4. Survival of daphnids in life‐table experiments decreased greatly after food became rare and was strongly reduced in those animals born during the clear‐water phase compared with those born later. In addition, age at first reproduction was retarded during the clear‐water phase, resulting in very low population growth rates. Survivorship patterns in life‐table experiments suggest a strong impact of non‐consumptive mortality on Daphnia population dynamics. 5. Field data of mortality point to differences in mortality patterns between years, probably resulting from different predation impacts of juvenile fish. In both years, however, adult mortality contributed substantially to overall mortality at the end of the clear‐water phase. As bottom‐up effects on D. galeata were very similar in both years, the significance of non‐consumptive mortality on the initiation of midsummer declines appears to depend largely on recruitment patterns before the clear‐water phase. A high impact can be expected when Daphnia populations are dominated by a peak cohort of nearly identical age during the clear‐water phase.     

Specificity of predator-induced neck spine and alteration in life history traits in Daphnia pulex

It has been proposed that the predator-induced defensive neck spine in Daphnia pulex has a demographic cost. Our results show that this cost is not merely an allocation cost related to the formation and maintenance of the neck spine. In a life table experiment, we tested whether spine induction and life history traits in D. pulex are affected by different invertebrate predators: first and third instar Chaoborus, fourth instar Mochlonyx and two size classes of Notonecta and Dytiscus larvae. D. pulex showed sensitivity to the different predators. Predator-exposure affected one or more of the following life history traits of D. pulex: the timing of first reproduction, clutch size, and growth. In some cases, exposure to predators altered life history traits when neck spine induction did not occur. These shifts in life history traits occurring in the absence of spine induction may be caused by behavioral or physiological changes triggered by the predators.     

Life history elasticity and the population-level effect of p-nonylphenol on Daphnia galeata

In order to evaluate population-level effects of p-nonylphenol on a cladoceran zooplankton (Daphnia galeata), the chronic effects on survival and reproduction were estimated with partial life table tests, which examined responses in life history characters until 3 weeks after birth. The observed responses in survival and reproduction were converted to reductions of the intrinsic rate of natural increase r. The population level EC₅₀, which is defined as the exposure concentration that reduces r by 50%, was estimated as 16.1 g l^–1. In order to examine the extent to which the population-level effect in terms of r is influenced by extra mortality in nature, which is induced by predation, starvation, etc., sensitivity (elasticity) measures of the intrinsic rate of natural increase to reductions in age-specific survival and reproduction were calculated under hypothetical predation schemes. The sensitivities of the intrinsic rate to changes in survival and reproduction invariably decline rapidly after the onset of reproduction irrespective of predation schemes. This implies that partial life cycle tests until 21 days after birth can provide reliable estimates of the population-level effects.     

Foraging strategies and growth inhibition in five daphnids feeding on mixtures of a toxic cyanobacterium and a green alga

1. Laboratory experiments were used to study the feeding, growth and reproduction of five daphnids in mixtures of a toxic cyanobacterium, Microcystis aeruginosa, and a green alga, Scenedesmus acutus. The mixtures included 0%, 20%, 50%, 80% and 100% Microcystis with a total food concentration of 0.5 mg C L^??1 in each treatment. The feeding rate was measured after 1 and 24 h of acclimatization to the mixtures. 2. Toxic Microcystis inhibited feeding in all the species, but they exhibited an unexpected diversity and complexity in the pattern of feeding inhibition. Daphnia magna exhibited the strongest inhibition of feeding after 1 h of exposure to toxic food, but had substantially recovered after 24 h in the same mixtures. This pattern of inhibition and recovery may balance the benefits of reduced ingestion of toxin with the disadvantage of a reduced energy intake. 3. All five daphnids grew quickly in the Scenedesmus control, whereas growth and reproduction declined with an increasing proportion of the toxic alga in the diet. Daphnia pulicaria showed the least inhibition of growth and reproduction, D. pulex showed the strongest inhibition and the three remaining species exhibited intermediate sensitivity. 4. Estimates of gross growth efficiency (GGE; growth/ingestion) provided a means for discriminating between the effects of feeding inhibition and direct toxicity on zooplankton growth. Daphnia pulex exhibited a sharp decline in GGE, suggesting that growth inhibition was a result of both feeding inhibition and direct toxicity. In contrast, D. magna exhibited a nearly constant GGE, indicating that feeding inhibition accounted for its decline in growth. However, two Daphnia species (i.e. D. pulicaria and D. galeata) exhibited improbable increases in GGE with toxic cyanobacteria, suggesting that their feeding rates were underestimated. Growth assays with sensitive and resistant zooplankton species are proposed for testing the potential impacts toxic cyanobacteria in lakes.     

Predator-induced phenotypic plasticity in Daphnia pulex: uncoupling morphological defenses and life history shifts

Chemical cues from a predator Chaoborus sp. induce morphological defense (neck spine) and life history shifts (later reproduction, decreased fecundity but larger juvenile size) in the waterflea Daphnia pulex. These shifts have been interpreted either as costs of defense or as separate adaptation. In order to investigate if the life history shifts can be separated from the morphological defense, Daphnia pulex individuals were exposed to chemical cues from Chaoborus at different stages of life for variable periods. The daphnids that were exposed to Chaoborus started their reproduction later than the controls, although the differences were not statistically significant. Neck spine was induced only if daphnids were exposed to Chaoborus in an early stage of their life. Numbers of eggs produced were not affected by the different treatments, but egg mortality was higher in mothers exposed to Chaoborus. With these treatments it was possible to see neck spine induction without measurable life history changes or costs. On the other hand, irrespective of neck spine presence, the Chaoborus chemical(s) had an effect on Daphnia pulex mothers.Publication no 2159. Netherlands Institute of Ecology, Centre for LimnologyPublication no 2159. Netherlands Institute of Ecology, Centre for Limnology     

Effects of salinity on the life history and fitness of Daphnia magna: variability within and between populations

The life history traits of Daphnia magna were studied in laboratory experiments under freshwater and brackish (5 salinity) conditions. The variability of responses within and between populations was examined by comparing 11 clones from a brackish lake and 10 clones from a freshwater pond. Experimental clones were hatched from ephippia collected from the sediment and thus represent random samples of the clone banks of each population.Most clones with a high salinity tolerance were from the population of the brackish habitat, but some were also found in the freshwater population. Thus, freshwater populations appear to have the potential to invade brackish habitats. A proportion of clones from the brackish population had very low fitness (measured as e^r) under freshwater conditions. This unexpected result means that freshwater adaptation can be lost by the freshwater cladoceran Daphnia magna. The effects of unfavourable conditions on growth and reproduction varied among clones and were not correlated. This clonal variation in growth and reproduction indicates that the environmental sensitivities of these traits are independent. The pattern of fitness reaction norms showed no trade-off between fitness under brackish and under freshwater conditions for either population. Thus, euryhaline generalists should be favoured in habitats with salinity fluctuations between freshwater and brackish conditions.     

Positive selection of digestive proteases in Daphnia: A mechanism for local adaptation to cyanobacterial protease inhibitors

Due to the combined effects of global warming and eutrophication, the frequency of deleterious cyanobacterial blooms in freshwater ecosystems has increased. In line with this, local adaptation of the aquatic keystone herbivore Daphnia to cyanobacteria has received major attention. Besides microcystins, the most frequent cyanobacterial secondary metabolites in such blooms are protease inhibitors (PIs). Recently, it has been shown that a protease gene showed copy number variation between four D. magna populations that differed in tolerance to PIs. From that study, we chose two distinct populations of D. magna which had or had not coexisted with cyanobacteria in the past. By calculating F_ST values, we found that the two populations were genetically more distant in the protease loci than in neutral loci. Population genetic tests applied to the tolerant population revealed that positive selection was most probably acting on the gene loci of the digestive protease CT448 and CT802. We conclude that the selection of digestive proteases and subsequent reduction in copy number is the molecular basis of evolutionary changes leading to local adaptation to PIs.     

Interspecific variation in the use of carotenoid-based coloration in birds: diet, life history and phylogeny

Birds show striking interspecific variation in their use of carotenoid-based coloration. Theory predicts that the use of carotenoids for coloration is closely associated with the availability of carotenoids in the diet but, although this prediction has been supported in single-species studies and those using small numbers of closely related species, there have been no broad-scale quantitative tests of the link between carotenoid coloration and diet. Here we test for such a link using modern comparative methods, a database on 140 families of birds and two alternative avian phylogenies. We show that carotenoid pigmentation is more common in the bare parts (legs, bill and skin) than in plumage, and that yellow coloration is more common than red. We also show that there is no simple, general association between the availability of carotenoids in the diet and the overall use of carotenoid-based coloration. However, when we look at plumage coloration separately from bare part coloration, we find there is a robust and significant association between diet and plumage coloration, but not between diet and bare part coloration. Similarly, when we look at yellow and red plumage colours separately, we find that the association between diet and coloration is typically stronger for red coloration than it is for yellow coloration. Finally, when we build multivariate models to explain variation in each type of carotenoid-based coloration we find that a variety of life history and ecological factors are associated with different aspects of coloration, with dietary carotenoids only being a consistent significant factor in the case of variation in plumage. All of these results remain qualitatively unchanged irrespective of the phylogeny used in the analyses, although in some cases the precise life history and ecological variables included in the multivariate models do vary. Taken together, these results indicate that the predicted link between carotenoid coloration and diet is idiosyncratic rather than general, being strongest with respect to plumage colours and weakest for bare part coloration. We therefore suggest that, although the carotenoid-based bird plumage may a good model for diet-mediated signalling, the use of carotenoids in bare part pigmentation may have a very different functional basis and may be more strongly influenced by genetic and physiological mechanisms, which currently remain relatively understudied.     

Essential fatty acid content and the phosphorus to carbon ratio in cultured algae as indicators of food quality for Daphnia

1. A series of experiments examined growth of Daphnia magna on three algal diets ( Rhodomonas minuta , Scenedesmus acutus and Synechococcus sp.) at varying physiological states [nitrogen and phosphorus (P) limitations] to test whether variation in algal fatty acid and/or elemental composition can predict Daphnia growth.
2. These algae differed widely in their essential fatty acid (EFA) composition while phosphorus (P) or nitrogen limitation had only a small influence on their ω3-polyunsaturated fatty acid (PUFA) content.
3. Individually, algal ω3-PUFA content explained 69% of the variation in the normalised growth of D. magna, while algal phosphorus content explained 11% of the variation. Quantitative models for D. magna growth used both algal ω3-PUFA content and algal C : P ratio as food quality indices. Together, algal ω3-PUFA content and C : P ratio explained 70% of the variation in the normalised growth rate of D. magna .
4. Our results indicate that EFA influenced algal food quality much more strongly than P. The EFA and mineral P impacts appear to be independent.     

Adult diet affects the life history and host-killing behavior of a host-feeding parasitoid

Neochrysocharis formosa (Westwood), an important biocontrol agent of agromyzid leafminers worldwide, is a host-feeding, idiobiont parasitoid. Female wasps have three types of host-killing behaviors: reproductive (parasitism), non-reproductive host feeding (host feeding), and host stinging without oviposition or feeding (host stinging). In this study, we compared the life history and host-killing behaviors of female parasitoids under four adult diets: starvation, hosts only, hosts plus honey (10% w/v honey solution), and honey only. Furthermore, we analyzed the host-feeding and oviposition preferences of adult females in the hosts-only and hosts-plus-honey treatments. Female parasitoids feeding on hosts had significantly increased longevity, higher fecundity, more host-stinging events, and caused a higher total host mortality than parasitoids in the starvation treatment. The honey supplement significantly increased longevity, fecundity, host-stinging events, and total host mortality, as well as average daily fecundity, but did not alter host-feeding events, host-stinging events, or daily total host mortality. However, the honey supplement did reduce the number of daily host-feeding events and induced a shift toward oviposition. Finally, we found that the non-reproductive host killing caused by host feeding and host stinging enhanced the control potential of N. formosa. These results should contribute to a better understanding of the biocontrol efficiency of destructive host feeders.     

Adaptive responses by mouse fetus to a maternal HLE diet by downregulating SREBP1: a microarray- and bio-analytic-based study

Maternal diet has long been recognized as a significant factor affecting offspring development and health, but the target genes affected by a maternal high-lipid diet are currently unknown. In this study, the gene expression profile of neonatal mouse liver was analyzed using gene chips to identify genes with significant up- or downregulated expression levels due to maternal high-fat diet during gestation. Real-time PCR and Western blotting were used to measure key genes selected using microarray. Serum lipid, glucose, and insulin levels in adult offspring from dams fed with chow or a high-lipid diet were measured using commercial kits. Results indicate that the expression of genes involved in cholesterol and fatty acid synthesis were significantly inhibited, while the expression of genes involved in glycolysis were significantly decreased by maternal high-lipid diet during gestation. SREBP1 might be the key gene regulating genes involved in fatty acid, glucose, and cholesterol metabolism in response to a maternal high-fat diet.