Evidence for strong host clone-parasite species interactions in the Daphnia microparasite system

Abstract Organisms are often confronted with multiple enemy species. Defenses against different parasite species may be traded off against each other. However, if resistance is based on (potentially costly) general defense mechanisms, it may be positively correlated among parasites. In an experimental study, we confronted 19 clones from one Daphnia magna population with two bacterial and three microsporidian parasite species. All parasites were isolated from the same pond as the hosts. Host clones were specific in their susceptibility towards different parasite species, and parasite species were host-clone specific in their infectivity, spore production, and virulence, resulting in highly significant host-parasite interactions. Since the Daphnia 's resistance to different parasite species showed no obvious correlation, neither general defense mechanisms nor trade-offs in resistance explain our findings. None of the Daphnia clones were resistant to all parasite species, and the average level of resistance was quite similar among clones. This may reflect a cost of defense, so that the cumulative cost of being resistant to all parasite species might be too high.     

Accelerated molecular evolution in halophilic crustaceans

In contrast to the stable ionic composition of the oceans, inland waters show striking diversity, possessing salt concentrations varying from I mM to 5 M. Although species diversity is highest in fresh water, some lineages have colonized hypersaline environments where they encounter elevated levels of both ultraviolet (UV) radiation and osmotic stress. This study compares rates of evolution in halophilic and freshwater taxa for two groups of microcrustaceans, anostracans and daphniids, from Australia and North America. The results establish that halophilic species show consistent rate acceleration, involving elevated levels of both insertion/deletion events and of nucleotide substitutions. The elevated pace of molecular evolution does not appear to be linked to selection or to other agents that are known to influence the supply rate of mutations, such as UV exposure, generation length, or shifts in metabolic rate. However, variance in ionic strength, which is known to have potent effects on DNA-protein interactions as well as on the structural properties of DNA and proteins, might account for the lowered fidelity of DNA replication in life from hypersaline settings. Regardless of its cause, the consistent rate acceleration in halophiles suggests that past efforts to employ sequence divergences to date events, such as the age of asexual lineages in Artemia, have resulted in serious overestimates. More generally, the results indicate that coordinated shifts in rates of molecular evolution may occur in lineages exposed to extreme environmental conditions.     

Quantitative genetic variation in Daphnia: temporal changes in genetic architecture

Nonadditive genetic variation and genetic disequilibrium are two important factors that influence the evolutionary trajectory of natural populations. We assayed quantitative genetic variation in a temporary-pond-dwelling population of Daphnia pulex over a full season to examine the role of nonadditive genetic variation and genetic disequilibrium in determining the short-term evolutionary trajectory of a cyclic parthenogen. Quantitative traits were influenced by three factors: (1) clonal selection significantly changed the population mean phenotype during the course of the growing season; (2) sexual reproduction and recombination led to significant changes in life-history trait means and the levels of expressed genetic variation, implying the presence of substantial nonadditive genetic variation and genetic disequilibrium; and (3) Egg-bank effects were found to be an important component of the realized year-to-year change. Additionally, we examined the impact of genetic disequilibria induced by clonal selection on the genetic (co)variance structure with a common principal components model. Clonal selection caused significant changes in the (co)variance structure that were eliminated by a single bout of random mating, suggesting that a build-up of disequilibria was the primary source of changes in the (co)variance structure. The results of this study highlight the complexity of natural selection operating on populations that undergo alternating phases of sexual and asexual reproduction.     

Molecular genetic analysis of Daphnia in the English Lake District: species identity, hybridisation and resting egg banks

• 1 Taxonomic issues in many Daphnia species complexes are often confused by a high degree of phenotypic plasticity and by interspecific hybridisation. Here, we employ molecular genetic tools to confirm the species composition and incidence of hybridisation in extant and resting egg populations of Daphnia from Windermere and Esthwaite Water in the English Lake District.
• 2 A combination of species‐diagnostic allozymes and mtDNA, confirms that contemporary populations are dominated by a single species, D. galeata.
• 3 We present the first account of genetic characterisation of dated ‘resting’ eggs using microsatellites and mtDNA, employing PCR‐based DNA recovery, thus providing a temporal dimension to taxonomic patterns. Thirty years ago, two species were present in Esthwaite, D. galeata and D. hyalina, but Windermere populations were dominated by D. galeata only.
• 4 The use of PCR‐based mtDNA RFLP analysis as a species‐diagnostic tool, and microsatellites to monitor clonal diversity, provide a valuable approach for long‐term studies, especially in populations free from the complicating effect of frequent hybridisation. The detailed limnological records available for many large lakes, and associated changes in land‐use, pollutants and climate, combined with long‐term ephippial molecular genetic data, provide opportunities for exploring natural and anthropogenic impacts on genetic and community structure.

     

Species-specific N and P release rates in Daphnia

Bioavailable N and P release rates by juveniles and adults of three Daphnia taxa (D. hyalina, D. galeata and its interspecific hybrids D. hyalina × galeata) were measured to assess the effect of weight and interspecific differences on these rates in Daphnia. Immobilized Scenedesmus obliquus cells were used to estimate the release rates. The specific release rate of N varied between 5.19–5.71 g N mg C^-1 h^-1 for juveniles and 3.00–3.42 g N mg C^-1 h^-1 for adults. P excretion rate ranged between 1.93–2.37 g P mg C^-1 h^-1 for juveniles and 1.00–1.24 g P mg C^-1 h^-1 for adults. Our results show that the taxonomic affiliation of Daphnia individuals did not affect their N and P release rates.     

EFFECT OF GRAZING-ASSOCIATED INFOCHEMICALS ON GROWTH AND MORPHOLOGICAL DEVELOPMENT IN SCENEDESMUS ACUTUS (CHLOROPHYCEAE)

Populations of the phenotypically plastic alga Scenedesmus acutus Meyen were cultured in standard medium or in medium with filtered water from a Daphnia culture to examine the expression of ecomorphs in both water types. A rapid formation of four- and eight-celled coenobia was observed in the presence of Daphnia water, but not in standard medium. Moreover, cell dimensions were increased in the Daphnia water containing medium. Population growth rates were comparable in both water types; however, the carrying capacity appeared significantly lower in Daphnia water. Daphnia needed to feed on digestible food to produce the colony-inducing chemicals; medium from starved animals and Daphnia fed polystyrene beads appeared inactive. Neither suspensions of homogenate of Scenedesmus and Cryptomonas nor auxins affected colony formation or growth rate in S. acutus. The colony-inducing infochemicals are probably not constituents of the algae themselves; however, modification of algal constituents in the digestive system of daphniids may be involved.     

Variation in the neonate size of Daphnia pulex: the effects of predator exposure and clonal origin

Daphnia pulex clones originating from twelve small pond habitats were exposed to chemical cues from a size-selective predator, larvae of the phantom midge Chaoborus. Exposure delayed the onset of reproduction and increased the size at first reproduction. On the other hand, the neonates produced by these larger mothers were slightly smaller than the neonates produced by the smaller control mothers. In cladocerans, neonate size is usually positively correlated to the size of the mother. Thus exposure to Chaoborus kairomone apparently had direct effects on neonate size counterbalancing the maternal effects. Daphnia clones collected from Chaoborus-free and Chaoborus-rich environments exhibited different responses. In first adult instar, the clones from Chaoborus inhabited environments increased their offspring size under Chaoborus exposure whereas clones from Chaoborus-free environments did not. This may reflect clonal adaptation to the predation prehistory of their original habitat since larger neonates more quickly reach a size protected from the predator.     

Increasing turbidity significantly alters the diet of brown trout: a multi‐year longitudinal study

The stomach contents of adult brown trout Salmo trutta from Lake Sorell, Tasmania, were analysed during 6 years of increasing turbidity to follow changes in the diet associated with dramatic changes in water clarity. Brown trout were sampled from 1996, when turbidity was 26 NTU, to 2001 when turbidity was 141 NTU. The mean relative volume of stomach contents decreased progressively to 2001, by which time it was only one sixth of that in 1996, and the mean diversity of prey in stomachs decreased from an average of more than six species per stomach in 1996 to one species in 2001. The species composition of stomach contents shifted from domination by the phreatoicid isopod Colubotelson sp., to the galaxiid fish Galaxias auratus and the amphipod Austrochiltonia australis , and then the cladoceran Daphnia carinata . To give an indication of diet changes over a typical yearly cycle in the current turbid state of the lake, a sample was taken from each season from December 2000 to September 2001. Two basic diets were found during the year; brown trout specialized on D. carinata in summer and autumn, and G. auratus in winter and spring. Mean diversity of prey was less than two species per stomach in all samples from 2000 to 2001, except for the sample from spring 2001 when it was 2·2 species per stomach, and the mean relative volume of stomach contents was more than three times greater in winter than any other season. The ways in which high turbidity may have influenced the changes in the brown trout diet observed since 1996 and the patterns evident during the seasons of 2000–2001 are discussed.     

HOLARCTIC PHYLOGEOGRAPHY OF AN ASEXUAL SPECIES COMPLEX I. MITOCHONDRIAL DNA VARIATION IN ARCTIC DAPHNIA

Pleistocene glacial cycles undoubtedly altered the evolutionary trajectories of many taxa, yet few studies have examined the impact of such events on genetic differentiation and phylogeography at large geographic scales. Here we present the results of a circumarctic survey of mitochondrial DNA diversity in members of the Daphnia pulex complex. The analysis involved the survey of restriction site polymorphisms in a 2100-bp fragment of the NADH-4 (ND4) and NADH-5 (ND5) genes for 276 populations representing the two major groups (tenebrosa and pulicaria) in this complex across their Holarctic range. A comparison of the distribution patterns for seven clades in this complex revealed very clear phylogeographic structuring. Most notably, pulicaria group lineages were restricted primarily to the Nearctic, with some colonization of formerly glaciated portions of northern Europe. This group was not detected from vast expanses of northern Eurasia, including the Beringian glacial refuge. In contrast, tenebrosa group haplotypes showed considerable intercontinental divergence between Eurasian and North American lineages, but were absent from Greenland and Iceland, as well as the Canadian arctic archipelago. Dispersal in Eurasia was primarily in a westerly direction from Beringia, whereas dispersal in the Nearctic followed proglacial drainage patterns. Long-distance dispersal of certain lineages was observed in both groups, and variation in haplotype richness and nucleotide diversity allowed us to make inferences about the positioning of putative glacial refugia. Overall, the phylogeographic pattern of diversification in this arctic complex is characterized by the apparently unique postglacial histories for each clade, indicating that even closely allied taxa can respond independently to the allopatric effects of glacial cycles. This is in sharp contrast to other phylogeographic studies of species assemblages from more southern (unglaciated) latitudes, which are often characterized by concordant patterns.     

Interclonal variation in diel horizontal migration behaviour of the water flea <Emphasis Type="Italic">Daphnia magna</Emphasis>—searching for a signature of adaptive evolution

In shallow temperate lakes, zooplankton populations may exhibit diel horizontal migration (DHM) and move towards macrophytes during the day to avoid fish. Using a natural Daphnia magna population, we undertook an experimental investigation aimed to describe the genetic variation for DHM and to study whether an adaptive micro-evolutionary response occurred to changes in macrophyte coverage and fish predation pressure through time. Twenty-seven D. magna clones were hatched from ephippia in the sediment of shallow Lake Ring, Denmark. This lake was eutrophied during the 20th century and was subject to restoration measures in the 1970s. The DHM behaviour of the clones was observed both in the presence and absence of fish kairomone. Significant interclonal variation in DHM behaviour occurred in both treatments. To study the micro-evolutionary response of the Lake Ring D. magna population, two approaches were used. First, we compared the DHM behaviour of clones derived from ephippia collected at different depths. A comparison was conducted between clones resurrected from the period of eutrophication (1960–1980) and from the period of recovery (1986–2000). A significant treatment (presence and absence of fish kairomone) × period interaction effect was identified, suggesting a significant micro-evolutionary response for DHM behaviour. The D. magna clones exhibited a significantly stronger horizontal migration response during the period of eutrophication than in the recovery phase. Second, clonal means, representing the influence of the genotype on the trait, were correlated with environmental conditions (macrophyte cover, fish predation pressure and Secchi depth). The results of this analysis also suggest that a micro-evolutionary response by Daphnia has occurred in reaction to changes in fish predation pressure. In periods with high fish predation pressure, Daphnia migrated more strongly towards the plants. Guest editor: Piet Spaak Cladocera: Proceedings of the 7th International Symposium on Cladocera