Parasite-mediated selection in experimental metapopulations of Daphnia magna

In metapopulations, only a fraction of all local host populations may be infected with a given parasite species, and limited dispersal of parasites suggests that colonization of host populations by parasites may involve only a small number of parasite strains. Using hosts and parasites obtained from a natural metapopulation, we studied the evolutionary consequences of invasion by single strains of parasites in experimental populations of the cyclical parthenogen Daphnia magna. In two experiments, each spanning approximately one season, we monitored clone frequency changes in outdoor container populations consisting of 13 and 19 D. magna clones, respectively. The populations were either infected with single strains of the microsporidian parasites Octosporea bayeri or Ordospora colligata or left unparasitized. In both experiments, infection changed the representation of clones over time significantly, indicating parasite-mediated evolution in the experimental populations. Furthermore, the two parasite species changed clone frequencies differently, suggesting that the interaction between infection and competitive ability of the hosts was specific to the parasite species. Taken together, our results suggest that parasite strains that invade local host populations can lead to evolutionary changes in the genetic composition of the host population and that this change is parasite-species specific.     

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.     

Genotypic selection in Daphnia populations consisting of inbred sibships

The genetic basis of fitness reduction associated with inbreeding is still poorly understood. Here we use associations between allozyme genotypes and fitness to investigate the genetic basis of inbreeding depression in experimental outdoor populations of the water flea, Daphnia magna. In Daphnia, a phase of clonal reproduction follows hatching from sexually produced resting eggs, and changes in genotype frequencies during the clonal phase can be used to estimate fitness. Our experiment resembles natural colonization of ponds in that single clones colonize an empty pool, expand asexually and produce sexual offspring by selfing (sisters mate with their clonal brothers). These offspring diapause and form populations consisting of selfed sibships in the following spring. In 12 of 13 experimental populations, genotypes of selfed hatchlings after diapause conformed to Mendelian expectations. During the subsequent ca. 10 asexual generations, however, genotype frequencies changed significantly at 19 of 27 single loci studied within populations, mostly in favour of heterozygotes, with heterozygosity at multiple loci affecting the change in genotype frequency multiplicatively. Because variance in heterozygosity among siblings at a given marker reflects only heterozygosity in the chromosomal region around this marker, our results suggest that selection at fitness-associated loci in the chromosomal regions near the markers were responsible for these changes. The genotype frequency changes were more consistent with selection acting on linked loci than on the allozymes themselves. Taken together, the evidence for abundant selection in the chromosomal regions of the markers and the fact that changes in genotype frequencies became apparent only after several generations of clonal selection, point to a genetic load consisting of many alleles of small or intermediate effects, which is consistent with the strong genetic differentiation and repeated genetic bottlenecks in the metapopulation from which the animals for this study were obtained.     

Phylogeography of Daphnia magna in Europe

The phylogeographical structure of the water flea Daphnia magna in Europe was analysed using a 609-bp fragment of the cytochrome oxidase subunit I gene. Only moderate overall genetic divergence was detected within Europe. We detected four genetically and, to a large extent, geographically distinct phylogroups within Europe. Our results suggest that these groups recolonized large parts of Europe around 100,000 BP from different refugia. Overall, the pattern suggests a high degree of provincialism with a patchy occurrence of specific lineages, thus confirming the highly subdivided genetic structure usually observed in freshwater zooplankton populations. Although the region around the Mediterranean Sea was only sampled patchily, we obtained strong indications for the occurrence of more divergent genetic lineages in this region. Comparing our European samples to samples from North America and Japan revealed a higher level of differentiation, reflecting limited intercontinental dispersal.     

温度和种间竞争对大型溞种群动态和两性生殖的影响

在20℃、25℃下,将大型溞和老年低额溞分别按7+3(B组),5+5(C组),3+7(D组)的组合进行混合培养,以及用单种培养(10+0(A组),0+10(E组))作为对照,研究了温度和种间竞争对大型溞种群动态和两性生殖的影响.实验结果表明:在混合培养时,大型溞对老年低额溞产生明显的竞争优势.20℃、25℃下,单种培养的老年低额溞最大种群密度分别为大型溞的2.31和1.97,而在混合培养下老年低额溞的种群密度明显低于大型溞,在实验25d后几乎全部死亡.25℃下两种溞的种群密度之间存在极显著的负相关性(C组:r=-0.508,n=30,P<0.01;D组:r=-0.483,n=30,P<0.01).在20℃、单种培养下,大型溞在首次产幼溞时即出现雄体,且种群密度与雄体密度呈显著的相关性(r=0.678,n=24,P<0.01).大型溞的最大雄体密度(106 ind.(200ml)~(-1))和最大雄体比例(36.8%)均出现在20℃、单种培养下.25℃下,大型溞在混合培养的B组和C组首次产幼溞时即出现雄体,且雄体在混合培养B组的比例达28.2%.大型溞在25℃、单种培养下没有产生卵鞍,在混合培养下总计产生66个卵鞍,其中空卵鞍占51.5%,而在20℃、混合培养下没有卵鞍产生.实验结果暗示:在较高的温度下,种间竞争刺激了大型溞雄体的产生和卵鞍的形成,高密度的雄体有助于大型溞孤雌生殖雌体向两性生殖雌体的转化.     

The effects of parasitism and inbreeding on the competitive ability in Daphnia magna: evidence for synergistic epistasis

Synergistic epistasis for fitness is often assumed in models of how selection acts on the frequency and distribution of deleterious mutations. Evidence for synergistic epistasis would exist if the logarithm of fitness declines more quickly with number of deleterious mutations, than predicted by a linear decline. This can be studied indirectly by quantifying the effect of different levels of inbreeding on fitness. Here, six sets (different genetic backgrounds) of three increasingly inbred Daphnia magna clones were used to assess their relative fitness according to changes in frequency in a competition experiment against a tester clone. A novelty of the mating procedure was that the inbreeding coefficients (F) of the three clones belonging to each set increased in steps of 0.25 independent of the (unknown) inbreeding coefficient of the common ancestor. The equal increase of the inbreeding coefficients is important, because deviations influence the quantification of inbreeding depression, its variance and the detection of epistasis. In a simple mathematical model we show that when working with a partially inbred population inbreeding depression is underestimated, the variance of fitness is increased, and the detection of epistasis more difficult. Further, to examine whether an interaction between inbreeding and parasitism exists, each inbred clone was tested with and without a microsporidium infection (Octosporea bayeri). We found a nonlinear decrease of the logarithm of fitness across the three levels of inbreeding, indicating synergistic epistasis. The interaction term between parasitism and inbreeding was not significant. Our results suggest that deleterious mutations may be purged effectively once the level of inbreeding is high, but that parasitism seems not to influence this effect.     

Genetic diversity of Daphnia magna populations enhances resistance to parasites

The diversity-disease hypothesis states that decreased genetic diversity in host populations increases the incidence of diseases caused by pathogens (= monoculture effect) and eventually influences ecosystem functioning. The monoculture effect is well-known from crop studies and may be partially specific to the artificial situation in agriculture. The effect received little attention in animal populations of different diversities. Compared with plants, animals are mobile and exhibiting social interactions. We followed the spread of a microsporidian parasite in semi-natural outdoor Daphnia magna populations of low and high genetic diversity. We used randomly selected, naturally occurring host genotypes. Host populations of low diversity were initially monoclonal, while the host populations of high diversity started with 10 genotypes per replicate. We found that the parasite spread significantly better in host populations of low diversity compared with host populations of high diversity, independent of parasite diversity. The difference was visible over a 3-year period. Host genotypic diversity did not affect host population density. Our experiment demonstrated a monoculture effect in independently replicated semi-natural zooplankton populations, indicating that the monoculture effect may be relevant beyond agriculture.     

The response of European Daphnia magna Straus and Australian Daphnia carinata King to changes in geomagnetic field

This study investigates the effects of lifelong exposure to reversed geomagnetic and zero geomagnetic fields (the latter means absence of geomagnetic field) on the life history of Daphnia carinata King from Australia and Daphnia magna Straus from Europe. Considerable deviation in the geomagnetic field from the usual strength, leads to a decrease in daphnia size and life span. Reduced brood sizes and increased body length of neonates are observed in D. magna exposed to unusual magnetic background. The most apparent effects are induced by zero geomagnetic field in both species of Daphnia. A delay in the first reproduction in zero geomagnetic field is observed only in D. magna. No adaptive maternal effects to reversed geomagnetic field are found in a line of D. magna maintained in these magnetic conditions for eight generations. Integrally, the responses of D. magna to unusual geomagnetic conditions are more extensive than that in D. carinata. We suggest that the mechanism of the effects of geomagnetic field reversal on Daphnia may be related to differences in the pattern of distribution of the particles that have a magnetic moment, or to moving charged organic molecules owing to a change in combined outcome and orientation of the geomagnetic field and Earth's gravitational field. The possibility of modulation of self-oscillating processes with changes in geomagnetic field is also discussed.     

The use of inorganic substances to stimulate gut evacuation in Daphnia magna

Observations were done on the effect of inorganic substances on the gut evacuation process in Daphnia magna. Procedures which accelerate this process are described.     

1,8-二羟基蒽醌对大型溞的毒性效应

The study on the acute,sublethal and chronic toxicity of 1,8-dihydroxyanthraquinone (1,8-dihAQ) to Daphnia magna showed that the 48 h LC₅₀ was 0.37 mg稬^-1,and the feeding behavior of Daphnia magna was severely affected by the compound.When exposed to 0.2 mg稬^-1 of 1,8-dihAQ for 5 h,the filtration and ingestion rate of Daphnia magna was inhibited by 97%.Chronic toxicity test results indicated that the reproduction ability decreased dramatically after exposing to sublethal concentration of 1,8-dihAQ.It could be inferred that reproduction parameters and intrinsic rate of natural increase were the sensitive parameters in characterizing sublethal toxicity.The NOEC and LOEC values for reproduction parameters were also given.     

有毒和无毒微囊藻对大型溞生长和繁殖的影响

近年来,随着工农业的快速发展和人们生活水平提高,一些城市生活污水和工农业废水流入江河、湖泊等淡水水体中,加剧了水体的富营养化和污染,导致蓝藻过度生长繁殖而形成水华,其中最常见的是微囊藻水华[1,2].     

Thermal avoidance and preference in Daphnia magna

1. Water fleas (Daphnia magna) bred at 23°C were non-responsive to temperatures between 13 and 25°C.

2. At the lower (11°C) and upper limits (30°C) their klinokinetic avoidance behaviour showed a larger intraindividual than interindividual variation.

3. Thermal sensitivity for avoidance responses in D. magna was about 1.5°C.

4. For D. magna bred for one parthenogenetic generation at 14°C heat avoidance temperature was about 8°C lower, and cold avoidance temperature was about 1°C higher than in D. magna from 23°C.

5. In group experiments the animals showed some preference for the acclimation temperature.

6. Cold induced stenothermy and warm induced eurythermy in D. magna were related to the mode of reproduction.

Mechanisms of energy allocation to reproduction in the cladoceran Daphnia magna Straus

Understanding rules of resource allocation within individuals is helpful in explaining population dynamics. This is particularly the case under the conditions of resource limitation that are commonly experienced by zooplankton. Here, we evaluate assumptions underlying models of resource allocation in Daphnia and test the predictions of two models of response to starvation.
We also test the predictions of two simple models concerning the mechanisms of egg provisioning in Daphnia magna. Our results: (1) demonstrate that provisioning is discontinuous, occurring over a discrete period of the instar; (2) support the hypothesis that egg production occurs in a serial fashion; (3) show that Daphnia magna responds to starvation by ceasing egg production but that there is an increase in mean size of eggs provisioned during the instar, prior to starvation; and (4) broadly support predictions that the response of Daphnia to starvation is an instantaneous cessation of reproduction but that there is a time lag before death.     

不同种类农药表面活性剂对大型溞的急性毒性

运用评价化学品对水生生物毒性的标准试验方法,比较了39种非离子型、6种阴离子型和3种阳离子型的常用代表性表面活性剂对大型溞的急性毒性.结果表明:3种阳离子型表面活性剂1427、1227及C8-10的急性毒性均为剧毒,其中1427毒性最高,EC50值为0.97×10-2 mg·L-1;非离子型表面活性剂中蓖麻油聚氧乙烯醚、吐温和斯潘系列乳化剂均为低毒,而烷基酚聚氧乙烯醚系列和脂肪醇聚氧乙烯醚系列表面活性剂的毒性稍偏高,而AEO-7和AEO-5的毒性达到高毒水平,EC50值分别为0.82和0.97 mg·L-1,且此类表面活性剂脂溶性越大,对大型溞的毒性越大;大部分阴离子表面活性剂的毒性为中毒,但NNO表现为高毒,EC50值为0.17 mg·L-1.     

The use of biomarkers in Daphnia magna toxicity testing

We investigated the effect of short-term exposure to cadmium and 2,4-dichlorophenoxy acetic acid on the digestive physiology of Daphnia magna and the consequences for the bioenergetics of the organism. In both cases, ingestion was more drastically reduced compared to digestive enzyme activity. Furthermore a differential shift in catabolism was noted: in general polysaccharidases were less affected than the enzymes responsible for protein and lipid digestion. Comparison of the ‘1 h in vivo fluorescence’ criterion (Janssen & Persoone, 1993) with the ingestion and digestive enzyme activity revealed that this rapid screening assay should be considered as a quantification of ingestion inhibition rather than a methodology assessing digestive enzyme inhibition. This revised version was published online in July 2006 with corrections to the Cover Date.     

温度和食物浓度对大型溞Daphnia magna种群动态和两性生殖的影响

在4个温度(15、20、25、30℃)和3个食物浓度(1×104,1×105,5×105 cells ml-1)下研究了大型溞(Daphnia magna)淮河种群的动态和两性生殖,结果表明:(1)在实验初期,大型溞种群密度不断增加,达到最大值后种群密度呈现缓慢下降或趋于稳定的趋势.除15℃下最大瞬时增长率(1.36 d-1)出现在中食物浓度组外,20℃(1.51 d-1)、25℃(1.39 d-1)、30℃(0.69 d-1)下的最大瞬时增长率均出现在高食物浓度组.(2)相同温度下,食物浓度与大型溞最大种群密度间存在显著正相关(p<0.01).(3)相同食物浓度下,随温度升高,大型溞首次产幼溞时间(5～21d)和到达最大种群密度的时间(15～29d)缩短,首次抱卵时的体长(1.75～2.67 mm)减少.(4)在较低的温度和中食物浓度组下,大型溞产生了较多的休眠卵或卵鞍(15℃:(16.7±2.5) ind. 和20℃:(18.3±3.8) ind.);30℃下没有出现休眠卵或卵鞍.除15℃、中食物浓度组外,其它实验组均产生雄体.实验25d后,雄体密度与种群密度呈显著的相关性(30℃:p<0.05;15～25℃:p<0.01).(5)本研究结果暗示大型溞休眠卵的形成受温度、食物浓度和种群密度的共同影响,且较低的温度是大型溞休眠卵形成的主要诱因.     

Sex ratio of the cyclic parthenogen Daphnia magna in a variable environment

The relationship between environmental factors, sex ratio and mating system in Daphnia magna was examined, and the adaptiveness of environmental sex determination over ametic sex determination was explored. Monthly sexual sex ratios (males over total number of males and sexual females) ranged from 0.31 to 1.0, the three-year average equalling 0.61. However, if only the samples collected during the period of frequent sexuality from August to October are included, the sexual sex ratio becomes equal, 0.51. Sexual sex ratios varied real between samples during the same period and the standard errors appeared highest in July ad August. Typical of suck times is some uncertainty in the environment, and the environmental cues can be contradictory. Sex expression in Daphnia appears to be determined by responses to complicated interactions between different environmental factors, which adaptively alter the sex ratio. The longterm sexual sex ratio of Daphnia aproaching the equilibrium 1:1, despite environmental sex %termination, gives support for Fisier's classic theory of equal parental investment in both sexes. An equal sex ratio is advantageous also during periods of small population size because it maximizes the effective population size.     

五氯酚对大型溞的急性亚慢性和慢性毒性

本文采用换水式试验研究了五氨酚溞（ＰＣＰ）对大型（Ｄａｐｈｎｉａｍａｇｎａ）的急性、亚慢性和慢性毒性,稀释水硬度为８０─１００ｍｇ／Ｌ（以ＣａＣＯ３计）。急性和慢性试验均使用小于一日龄的幼溞,试验温度为２５─２６℃,慢性试验进行了２０ｄ。用小于一日龄幼溞进行的亚慢性试验暴露了１９ｄ,而用四日龄幼溞的亚慢性试验则进行了１６ｄ,水温均保持在１９─２０℃。ＰＣＰ对大型溞的２４ｈ和４８ｈＥＣ５０分别是４８９和２４５μｇ／Ｌ。依据第１胎所产幼溞数求得的最低可观察效应浓度（ＬＯＥＣ）和无可观察效应浓度（ＮＯＥＣ）,在慢性试验中分别是１６０和８０μｇ／Ｌ,在１９ｄ亚慢性试验中分别为２００和１００μｇ／Ｌ,二者相近。试验结果表明,第１胎所产幼溞数是敏感的指标。     

Extinction hazards in experimental Daphnia magna populations: effects of genotype diversity and environmental variation

Extinction is ubiquitous in natural systems and the ultimate fate of all biological populations. However, the factors that contribute to population extinction are still poorly understood, particularly genetic diversity and composition. A laboratory experiment was conducted to examine the influences of environmental variation and genotype diversity on persistence in experimental Daphnia magna populations. Populations were initiated in two blocks with one, two, three, or six randomly selected and equally represented genotypes, fed and checked for extinction daily, and censused twice weekly over a period of 170 days. Our results show no evidence for an effect of the number of genotypes in a population on extinction hazard. Environmental variation had a strong effect on hazards in both experimental blocks, but the direction of the effect differed between blocks. In the first block, variable environments hastened extinction, while in the second block, hazards were reduced under variable food input. This occurred despite greater fluctuations in population size in variable environments in the second block of our experiment. Our results conflict with previous studies, where environmental variation consistently increased extinction risk. They are also at odds with previous studies in other systems that documented significant effects of genetic diversity on population persistence. We speculate that the lack of sexual reproduction, or the phenotypic similarity among our experimental lines, might underlie the lack of a significant effect of genotype diversity in our study.     

Coping with predator stress: interclonal differences in induction of heat-shock proteins in the water flea Daphnia magna

Although predation is a strong selection pressure, little is known about the molecular mechanisms to cope with predator stress. This is crucial to understanding of the mechanisms and constraints involved in the evolution of antipredator traits. We quantified the expression of heat-shock protein 60 (Hsp60), a potential marker for predator stress, in four clones of the water flea Daphnia magna, when exposed to fish kairomones. Expression of Hsp60 induction increased after 6 h and returned to base levels after 24 h of predator stress. This suggests that it is a costly transient mechanism to temporarily cope with novel predator stress, before other defences are induced. We found genetic variation in the fixed levels and in the fish-induced levels of Hsp60, which seemed to be linked to each clone's history of fish predation. Our data suggest that Hsp60 can be considered part of a multiple-trait antipredator defence strategy of Daphnia clones to cope with predator stress.