Morphological differentiation within the Daphnia longispina group

Although morphological evolution is assumed to be slow within Daphnia species complexes, discontinuities in morphological space can be detected. Here, morphological data derived from females of genetically-defined clones (cf. Gießler et al., 1999) are presented, in order to estimate the genetic component of phenotypic variance under standardised laboratory conditions. Animals originated from clonal assemblages of pre-alpine lakes and ponds, and a remote lake in western Germany, covering a wide range of morphotypes known from the traditional species D. cucullata, D. galeata, D. hyalina, D. rosea, and a variety of interspecific hybrids. Phenotypic analyses were based on quantitative and qualitative morphological characters of females in the first and fifth instars. Morphological divergence between clones was analysed using discriminant analysis or multidimensional scaling and the significance of the morphological groupings was estimated using neighbour-joining trees and bootstrapping. All analyses confirmed that (a) phenotypic similarities among taxa change with instar, (b) in contrast to low genetic divergence, pronounced morphological divergence exists between animals separated on the lake/pond level favouring speciation by the habitat shift hypothesis.     

A straightforward multiallelic significance test for the Hardy-Weinberg equilibrium law

Much forensic inference based upon DNA evidence is made assuming Hardy-Weinberg Equilibrium (HWE) for the genetic loci being used. Several statistical tests to detect and measure deviation from HWE have been devised, and their limitations become more obvious when testing for deviation within multiallelic DNA loci. The most popular methods-Chi-square and Likelihood-ratio tests-are based on asymptotic results and cannot guarantee a good performance in the presence of low frequency genotypes. Since the parameter space dimension increases at a quadratic rate on the number of alleles, some authors suggest applying sequential methods, where the multiallelic case is reformulated as a sequence of "biallelic" tests. However, in this approach it is not obvious how to assess the general evidence of the original hypothesis; nor is it clear how to establish the significance level for its acceptance/rejection. In this work, we introduce a straightforward method for the multiallelic HWE test, which overcomes the aforementioned issues of sequential methods. The core theory for the proposed method is given by the Full Bayesian Significance Test (FBST), an intuitive Bayesian approach which does not assign positive probabilities to zero measure sets when testing sharp hypotheses. We compare FBST performance to Chi-square, Likelihood-ratio and Markov chain tests, in three numerical experiments. The results suggest that FBST is a robust and high performance method for the HWE test, even in the presence of several alleles and small sample sizes.     

Hardy-Weinberg定律在专科教学中的课堂设计

Hardy-Weinberg定律是群体遗传学的第一理论基石,也是现代进化论、现代优生学和群体育种的理论基础,是遗传学教学中的重难点内容,但通过合理的教学设计可帮助学生全面理解、掌握并应用该定律,为后续学习奠定基础。     

Chaoborus and fish-mediated influences on Daphnia longispina population structure,dynamics and life history strategies

Summary This study examined the long term effects of predation by larvae of the midge Chaoborus and simulated fish predation on experimental Daphnia longispina populations. Chaoborus predation, relative to fish predation, led to populations composed of larger individuals as a whole, larger egg-bearing individuals, and a larger primiparous instar. Daphnia retained helmets beyond the first instar in response to the presence of Chaoborus. Both types of predation, relative to predator-free controls, reduced prey population size and rates of increase, but increased population death rates. The reduction in population size due to predation led to increased resource availability for individuals remaining in the populations and increased individual fecundity in the predation treatments. The differences noted between the Chaoborus, fish, and control treatments increased with predation intensity.     

Genetic variation in a subtropical population of Daphnia

We assayed a subtropical population of Daphnia ambigua for genetic variation using protein electrophoresis (9 loci) and quantitative genetics approaches (life history characters). Our goal was to obtain information about relative levels of variation in a subtropical population, and compare them with extensive previous studies in the temperate and arctic zones. The observed level of allozymic variation (H = 0.11) was consistent with those previously observed in other temperate zone Daphnia populations. However, variation for quantitative traits (heritability) was lower than typically observed in previously-studied temperate populations: estimates were not statistically different from zero. Because allozyme heterozygosity was consistent with previous temperate zone estimates, and the polymorphic allozyme loci did not depart from the expectations of Hardy-Weinberg equilibrium, we concluded that a period of clonal selection was the most likely explanation for the low heritabilities observed. We do not conclude that this study provides evidence to suggest that subtropical populations harbor lower levels of genetic variation because of their location.     

Plankton community of a polyhumic lake with and without Daphnia longispina (Cladocera)

The impact of Daphnia longispina (Cladocera) on the plankton food web was studied in a polyhumic lake where this species comprised almost all zooplankton biomass. Plastic enclosures (volume 7 m³) were inserted into the lake retaining the initial water stratification except that in one enclosure zooplankton was removed. After the removal of Daphniaa rotifer, Keratella cochlearis, ciliates and heterotrophic nanoflagellates increased markedly and the density and biomass of bacteria decreased. Edible algal species, Cryptomonas rostratiformisand three small chrysophytes,Ochromonas, Pedinella and Spinifermonas, took advantage of the removal of Daphnia, while more grazing-resistant species declined. In spite of the changes in the species composition of phytoplankton, the removal of Daphnia did not affect the biomass, primary production or respiration of plankton. The results implied that the density of heterotrophic flagellates and ciliates was controlled by Daphnia, but in its absence the former took its role as the bacterial grazers.     

Vertical distributions of bacteria and algae in a steeply stratified humic lake under high grazing pressure from Daphnia longispina

The vertical distributions of bacteria and algae in a steeply stratified, highly humic lake were studied during three 24 h periods in summer. The highest bacterial and algal densities and biomasses were recorded in the anoxic hypolimnion. The bacterial biomass in the hypolimnion was composed mainly of photosynthetic green sulphur bacteria (Chlorobium) which occurred at very low light intensity (< 1.5 μmol m⁻² s⁻¹). The numbers and biomasses of bacteria, both in the epilimnion and hypolimnion, were mostly higher at night than during the day, indicating possible asynchrony between the production and loss of bacteria. Because of vertical migration, the diurnal vertical distribution of algae was more variable than that of bacteria. Particularly in July and August, when cryptomonads were abundant, the biomass of algae was much higher in the epilimnion during the day than at night. The flagellated chlorophytes, Chlamydomonas spp. and Scourfieldia cordiformis, stayed mainly in the upper hypolimnion close to the oxic-anoxic boundary zone where only a small proportion of Daphnia longispina was continuously present. Unpalatable Mallomonas chrysophytes with silicified plates and bristles, and small, presumably heterotrophic, flagellates stayed in the oxic epilimnion together with a dense (up to 300 ind l⁻¹) population of D. longispina. The results indicated that, besides the physical and chemical properties of the water column, grazing pressure by Daphnia longispina strongly affected the vertical distribution of microorganisms in this polyhumic lake.     

Elemental composition of two coexisting Daphnia species during the seasonal course of population development in Lake Constance

Summary In 1986 and 1987 the seasonal course of population density, life history traits and biomass composition (CHN-elemental and ash) of two coexisting Daphnia species from L. Constance (Überlinger See) were studied. The two daphnids are morphologically similar, but demonstrate different seasonal variation of behaviour (i.e. diurnal vertical migration). With the exception of nitrogen content, the mean biomass composition of adults and that of eggs were significantly different: the dry biomass of eggs (adults) contained on average 55.6 (46.1) % C, 7.9 (6.5) % H, 9.7 (9.7) % N, and 10.0 (23.8) % ash. These values were similar to those of other daphnids; it may be concluded that the average daphnid composition is not species specific, but is determined mainly by the position in the life cycle and by the nutritional state of the animal. A distinct seasonal variation in the elemental composition of the biomass of adult daphnids was observed. It related significantly to the concentration of food, as indicated by the epilimnetic chlorophyll-a concentrations and by Secchi depth. From the amplitude of seasonal variation of elemental composition and from the shape of relationships between the elemental composition of daphnids and their food, it can be concluded that in summer the non-migrating D. galeata lives under stronger food limitation than the migrating H. hyalina. In spring, there were no differences in the biomass composition of either species. In summer, the adult fcmales of D. galeata contained significantly less carbon, and hydrogen than those of D. hyalina. In most parameters of biomass (i.e. biomass composition of adults) and of life history (i.e. clutch size, mean adult weight), D. galeata displayed higher coefficients of variation. A tendency in D. galeata to have peaks of population development carlier in the season and in D. hyalina to have them towards autumn was observed. These contrasting responses of the populations development, the seasonal variability of the biomass composition seem to support the view of Geller (1986) who called D. galeata and D. hyalina exploitative and conservative strategists respectively.This research was supported by the Deutsche Forschungsge-meinschaft within the Sonderforschungsbereich Cycling of Matter in Lake Constance (SFB 248), and by scholarship to R.B. from the Deutscher Akademischer Austauschdienst (DAAD)     

Population Dynamics of Daphnia Hyalina Leydig (Crustacea: Cladocera) in a Productive and an Unproductive Lake in the English Lake District

The population dynamics of Daphnia hyalina Leydig in a productive lake, Esthwaite Water, and an unproductive lake, Buttermere, in the English Lake District have been compared. The winter is passed as resting eggs in the bottom sediments in Buttermere and as free-swimming individuals in the planktonic zone in Esthwaite Water. In Esthwaite Water seasonal periodicity was characterised by maxima in spring and autumn and a minimum in summer; in Buttermere, there was no spring maximum and the first increase in population density was in summer. Population densities were higher and adult females were larger and laid more eggs per clutch in Esthwaite Water than in Buttermere. In each lake males became numerous in autumn. Observed rates of population increase, r and calculated birth rates, b and death rates d were nearly always higher in Esthwaite Water than in Buttermere.     

A new method of testing for Hardy-Weinberg equilibrium and ordering populations

The assumption of Hardy-Weinberg equilibrium (HWE) among alleles in a nonevolving population is of fundamental importance in genetic studies. Deviation from HWE in a population usually indicates inbreeding, stratification and sometimes problems in genotyping. In populations of affected individuals, these deviations can also provide evidence for association. In this paper, we introduce a measure based on the Kullback-Leibler discrimination information function that quantifies the deviation from HWE in a population. We use this measure to order populations. We also propose a test for HWE based on an estimate of this measure. The test is a statistically consistent test of the null hypothesis for all alternatives and is very easy to implement. Our proposed test statistic is compared with an earlier, widely used, test. Finally, the use of the proposed new test is shown in an illustrative example.     

Mechanisms of the filtering area adaptation in Daphnia

The changes of selected parameters of the filtering comb of the third thoracic limb were studied in a natural population of Daphnia pulicaria Forbes, as well as in experimental enclosures and in lab cultures, including individual life history. Two hypotheses were tested: 1. either these changes are related to the succession of clones coexisting within one population, or 2. the size of the filtering area changes gradually as an individual adaptation during the moulting. No evidence supporting the clonal hypothesis was found. On the contrary, the adaptability of the filtering comb is the same in a natural population as it is in a clone and in individuals.     

Diel vertical migration and feeding rhythm of Daphnia longispina and Bosmina corexoni in Lake Toya, Hokkaido, Japan

Diel vertical migration (DVM) and diel feeding rhythm of two cladocerans, Daphnia longispina and Bosmina coregoni were investigated at the pelagic area of Lake Toya (Hokkaido, Japan) in May, August and October 1992. Both species performed nocturnal DVM. The amplitude of DVM, however, became smaller from May to October. Such seasonal variations in DVM could not be explained by light penetration and/or water temperature. The two species had a clear feeding rhythm; they fed at night in May and October but also after sunrise in August. These feeding rhythms appeared to be related to the light-dark cycle, but were not necessarily associated with their DVM. We suggest that the diel feeding rhythm and DVM are regulated independently by light cues.     

Seasonal dynamics of metabolic activity of the Daphnia community in Lake Bled (Slovenia)

Seasonal changes of the community composition, oxygen consumption (R) and respiratory electron transport system (ETS)-activity of the Daphnia community living in Lake Bled (Slovenia) were studied between January and December 1998. The ETS activity of ovigerous Daphnia females at in situ temperature ranged from 3.27 l O₂ mg dw^–1 h^–1 in February to 20.91 l O₂ mg dw^–1 h^–1 in July. Respiration rates at in situ temperature varied from 4.04 l O₂ mg dw^–1h^–1 in December to 18.68 l O₂ mg dw^–1 h^–1in July. The influence of four factors (temperature, body size, fecundity, genetic differences) on the metabolism were investigated. Both ETS activity and respiration rate significantly correlated with temperature. The proportion of hybrid D. cucullata× galeata in Daphnia community correlated significantly with respiration rate at in situtemperature and ETS activity at standard temperature also. ETS activity and respiration rate showed no significant correlation with body size and the fecundity of Daphnia, whereas ETS activity in D. hyalina × galeata and D. cucullata× galeataseparately correlated with body size. ETS activity of D. hyalina × galeata also was correlated with fecundity. Hybrid D. hyalina× galeata had up to one third lower ETS activity than D. cucullata× galeata. The mean ETS/R ratio in the Daphnia community was 1.16±0.28 (N= 12). The ETS/R ratio did not correlate significantly with temperature, body size, fecundity or the proportion of D. cucullata× galeatain the Daphnia community. Laboratory experiments showed that both hybrids had similar ETS/R ratios.     

A note about Daphnia in Venezuelan waterbodies

After examining numerous water samples of 19 lakes and reservoirs in Venezuela, only two species of Daphnia were found: D. laevis (in colder, high altitude lakes) and D. gessneri. This observation supports the theory of the scarcity of Daphnia species in tropical inland waters.     

Seasonality,Population Dynamics and Production of Daphnia longispina in the Subtropical Lake Bhimtal (U.P.), India

Seasonal variation in population density, birth rate, mortality and production of a daphnid was studied in a high, altitude sub-tropical lake. A bimodal growth curve with maxima during summer and autumn was observed. A positive correlation was reported between fecundity and death rate, probably due to a difference in age specific death rate combined with a high relative frequency of juveniles in a growing population. The production of Daphnia longispina correlates better with temperature than with food. The P/B ratio was higher during summer and autumn months.     

Intersex and male development in Daphnia magna

Sex induction is environmentally stimulated in Daphnia and involves a cue-dependent response for sex determination. Somatic growth was shown to be similar in males and females during juvenile instars, but divergent due to a reduction in male somatic growth, at about the time that females produce ovaries. At this time, males appeared morphologically adult with respect to secondary sex characteristics. Intersex was rare and is unlikely to be important in natural populations. Intersex could be induced in both sexes, and observed more frequently after longer exposure to high temperature, or in the second generation following a temperature change. This indicates an impact on the ability of mothers to determine the sex of the offspring. It may be possible to use intersex characteristics for manipulative investigation of sex-determination mechanisms in Daphnia. Although sex-determination is initiated before birth, intersex occurrence suggests that development of male characters requires an additional process, probably involving hormone activity, during juvenile development and maturation.     

Competition in natural populations of Daphnia

I investigated the competitive relationships between two species of Daphnia, D. galeata and D. cucullata, and their interspecific hybrid. The term hemispecific competition was introduced to describe competition between parental species and hybrids. In eutrophic Tjeukemeer both parental species were found to compete with the hybrid, whereas competition between D. galeata and D. cucullata seemed limited. Although the effect of competition on life history traits of daphnids may be profound, the influence of the competitors on the seasonal dynamics of the Daphnia species seems limited.     

Daphnia vertical distribution and the presence of toxic cyanobacteria

Daphnia can alter its vertical position in the water column in response to chemical cues from predators. In this study we tested the hypothesis that a Daphnia pulex clone with little evolutionary exposure to cyanobacteria would move away from patches of cyanobacteria (Anabaena affinis and A. flos-aquae) which contain potent endotoxins. Daphnia was censused at 2 h intervals for 6 h in laboratory columns in which there was a steep vertical gradient of cyanobacteria. Data were analyzed by repeated-measures ANOVA. Control (no Anabaena) and experimental columns showed no significant differences in Daphnia distributions. Daphnia in experimental columns frequently moved into areas with dense concentrations of Anabaena and stayed there for long periods of time. Our results show that this D. pulex clone does not exhibit a rapid (within 6 h) avoidance response to toxic Anabaena.     

Daphnia exhibit diurnal vertical migration in shallow rock-pools

We report diurnal changes in vertical distribution ofDaphnia longispina andD. magna in shallow (25–60 cm) rock-pools on Baltic islands, southern Finland. During the daytime shallow rock-poolDaphnia reside in water layers close to the sediment while at midnight they either are evenly distributed throughout the water column, or have a modal peak of distribution in the surface water. Contrary to this pattern, no changes were observed in vertical distribution ofD. longispina at noon and at midnight in a 150 cm deep pond. We suggest that the pattern ofDaphina vertical migration in shallow rock-pools is not satisfactorily explained by predation.