Fitness and sexual response to population density in Daphnia pulex

1. The switch between asexual and sexual reproduction is an important fitness component in cyclically parthenogenetic populations as it is the key to persistence in unstable habitats and because it influences population genetic characteristics such as linkage disequilibrium and population genetic structure.
2. Genetic variation for sexual and asexual reproductive rate ( R _C) was examined, under varying population density, in Daphnia pulex sampled from a rockpool system and two other distinct European localities.
3. Density affected the switch to sexual behaviour, as found in other studies and reduced R _c. Production of males was not correlated with sexual reproduction, promoting outcrossing.
4. Genetic variation was found for the response to density, both within and between populations. There was substantial variation within the rockpools despite their demographic instability, indicating rapid recovery of life-history variation or maintenance during bottlenecks.     

Direct distributional response in Daphnia pulex to a predator kairomone

The hypothesis that zooplankton are capable of direct and differentiatedbehavioural responses to different biotic factors was tested.A multichamber flow-through technique enabled predeterminedfood levels of the green alga Scenedesmus acutus to be maintainedconstant throughout the experiments. By adding water from aculture of fourth instar Chaoborus flavicans larvae to one ofthe end chambers, a concentration gradient of the chemical substancewas established across the parallel chambers. Evidence is presentedfor a direct avoidance response in the water flea Daphnia pulexto a substance released by the predatory midge C.flavicans.This response occurs in the absence of visual cues and is independentof food availability. Finally, the results indicate that swarmingin D.pulex may occur in the absence of both light and predatorscent, but is influenced by food concentrations.     

Inter-genotype variation in reproductive response to crowding among Daphnia pulex

Crowding is known to have a major influence on reproduction in the freshwater microcrustacean Daphnia pulex. We analyzed reproductive output of six different D. pulex genotypes under two different density regimes in the laboratory. Four of these genotypes reproduce via obligate parthenogenesis, allowing thorough analysis of the life history strategies of some asexual lines. Among 30,109 neonate offspring and 1041 resting egg ephippia collected, several trends were evident. Crowding induced increased resting egg production and reduced neonate offspring production among all genotypes. Offspring sex ratios grew more male-biased with maternal age. The extent, but not direction, of each of these trends varied among genotypes. Offspring sex ratios, and the very direction in which they changed in response to crowding, differed significantly among genotypes with some genotypes producing more and others fewer males in response to crowding. Obligately parthenogenetic genotypes seemed to respond to the crowding stimulus in similar ways as the facultatively parthenogenetic genotypes, as expected from the sexual origins of their genomes. The inter-genotype variation in life-history traits observed in this and other investigations calls into question the common practice of extrapolating results from a single Daphnia genotype to an entire species. Our findings are considered in the context of other research in the field of environmental influences on Daphnia reproduction with a review of representative literature.     

Depth selection by Daphnia pulex in response to Chaoborus kairomone

1. The presence of kairomone from the predaceous larval dipteran Chaoborus americanus can result in juvenile Daphnia pulex positioning themselves higher in the water column. Chaoborus normally lives at greater depth during the day, so this behavioural response by Daphnia will reduce its encounter rate with the predator and enhance survival. 2. Behavioural observations of seven clones from a single population were made under four treatments (with and without Chaoborus kairomone prior to and/or during experiments). 3. All clones moved higher when exposed to kairomones during behavioural observations. Six moved higher when kairomone was added prior to the experiment (i.e. pre‐conditioned), while the other clone went lower, providing evidence for the presence of genetic variation in induced behaviour. When this clone was removed from the analysis, the evidence for genetic variation in induced response disappeared. 4. Juvenile Daphnia that were pre‐conditioned had a significantly greater response than those that had no previous exposure. Of the total shift in depth (comparing treatment and control means), 38% was due to prior exposure to the kairomone (‘preconditioning’), while 62% was due to exposure during the 2‐h experiments when the depth selection was assessed. When the effect of the one clone with a qualitatively different response was removed, these two figures became 45% and 55%, respectively. 5. The enhanced effect of prior exposure to the kairomone during development suggests that such exposure causes greater sensitivity to the kairomone.     

Horizontal distribution of Daphnia pulex in response to toxic and non-toxic algal extracts

We studied the horizontal distribution of Daphnia pulex as aresponse to gradients of toxic extracts of two cyanobacteria,Microcystis aeruginosa and Oscillatoria agardhii and non-toxicextracts of the green alga Scenedesmus obtusiusculus. The experimentwas performed in the dark using a flow through system with fivechambers. Daphnia pulex aggregated at the inlet of Scenedesmusextract, but showed similar responses to extracts of toxic cyanobactenaand food-free medium. Chemical cues seem to affect food search,but do not effect Daphnia pulex's avoidance of toxic algae.     

Intraspecific variability in response to stimuli for male and ephippia formation in Daphnia pulex

Natural clones and clones from laboratory crossings of Daphnia pulex have been tested in their response to stimuli for male and ephippia formation. The clones are from ponds in Illinois, USA, an area with both obligate and cyclical clones of D. pulex. The progeny from two types of crosses were studied: 1) crosses between an obligate clone producing males and a cyclical clone producing only females, and 2) self-fertilization within a cyclical clone producing both sexes. Both the natural and the artificial clones showed great variation in response to environmental stimuli for males. Ephippia response seems more equal among the clones, but two of the artificial clones could hardly be stimulated to produce ephippia in the induction experiments. In crowded cultures, however, they produced a few. Crossings between cyclicals and obligates yielded mostly cyclical progeny. Progeny from the selfed clone had low survival and fecundity.     

Demographic costs of Chaoborus-induced defences in Daphnia pulex: a sensitivity analysis

Summary We examined the demographic costs of Chaoborus-induced defensive spine structures in Daphnia pulex. Our aim was to assess the role of resource limitation and the interaction effects of limiting food level and antipredator structures on fitness of D. pulex and to pinpoint those life stages that are most sensitive to changes in the defence regime. Chaoborus-induced and typical morphotypes of D. pulex were reared at high and low food concentrations. Instar-based matrix population models were used to quantify the effects of predator-induction, food and their interaction on fitness of D. pulex. Predator-induction caused a statistically significant reduction in fitness at low food levels, but not at high food levels. Sensitivity analyses revealed that the fitness effects were primarily due to changes in the growth rate in instars 1–5, and secondarily to small reductions in the fertility of instars 5–10. The interaction between Chaoborus exposure and low food concentration was negative, and mediated through growth and fertility components. Both these components were reduced more in the Chaoborus-exposed, low food treatment than would be expected in the absence of interaction.     

Physiological responses of Daphnia pulex to acid stress

Background  

Acidity exerts a determining influence on the composition and diversity of freshwater faunas. While the physiological implications of freshwater acidification have been intensively studied in teleost fish and crayfish, much less is known about the acid-stress physiology of ecologically important groups such as cladoceran zooplankton. This study analyzed the extracellular acid-base state and CO₂ partial pressure (P _CO2), circulation and ventilation, as well as the respiration rate of Daphnia pulex acclimated to acidic (pH 5.5 and 6.0) and circumneutral (pH 7.8) conditions.     

Reproducibility of short-term and reproduction toxicity experiments with Daphnia magna and comparison of the sensitivity of Daphnia magna with Daphnia pulex and Daphnia cucullata in short-term experiments

Short-term toxicity tests with Daphnia magna are reasonably reproducible. The sensitivity of Daphnia pulex, Daphnia cucullata and Daphnia magna proved to be about the same in short-term tests with 15 different chemical compounds.Reproduction toxicity experiments with Daphnia magna can easily be carried out within three weeks. At least duplicate experiments are necessary to arrive at an approximate no-effect level.     

Embryology of Chaoborus-induced spines in Daphnia pulex

Daphnia pulex (Crustacea: Cladocera) embryos were found to be sensitive to a chemical cue (kairomone) in an extract of the predator Chaoborus americanus (Insecta:Diptera). Sensitivity of embryos to the kairomone remains throughout embryonic development. Apparently declining sensitivity as development proceeds may be due to the amount of time the embryos are exposed to the kairomone. Male embryos were also found to be sensitive to the kairomone. The smallest eggs within a brood produced small offspring, which showed the antipredator morphology to a significantly lower degree than largest eggs. The production of the neckteeth is described, at the developmental stage in the maturation of the Daphnia coinciding approximately with the escape of the embryos from the brood chamber.     

The cost of males in Daphnia pulex

Sex is paradoxical, because asexuals should replace their sexual ancestors by avoiding the demographic cost of producing males (hereafter referred to as the cost‐of‐males). Despite the large body of theoretical and empirical work dealing with the paradox of sex, the cost‐of‐males assumption has been rarely tested. In the present study, we tested the cost‐of‐males assumption in the cladoceran Daphnia pulex. Populations of this species consist of both cyclically parthenogenetic (i.e. sexuals) and obligately parthenogenetic (i.e. asexuals) lineages. In addition, some of the asexual lineages produce only female offspring, whereas others produce functional males, which can mate with sexual females. We compared the reproductive investment of sexuals, male‐producing asexuals, and non‐male‐producing asexuals when raised separately under various environmental conditions. We also determined the outcome of competition between pair‐wise combinations of these reproductive modes. The cost of males was evident when sexual and asexual females were raised separately: sexuals produced fewer female offspring. However, there was no cost of males when reproductive modes were raised in pairs, as sexuals won the competition with asexuals. Our results directly relate to the field conditions experienced by D. pulex. Sexuals might suffer the cost of males at the beginning of the season, when resource competition is low; but when conditions deteriorate as the population approaches carrying capacity, sexuals seem to be better competitors in spite of male production.     

A comparison of the sensitivities of Daphnia magna and Daphnia pulex to six different cyanobacteria

Cyanobacteria have become an important environmental concern due to their ability to produce a wide range of natural toxins. At present, very few studies describe concentration response curves for cyanobacteria other than Microcystis. However, field evidence highlights that both cyanobacterial concentration as well as cyanobacterial species composition vary considerably with season and year. Therefore, the aim of this study was to investigate the effects of different cyanobacteria at various concentrations of these cyanobacteria in the diet on the reproduction of Daphnia pulex and Daphnia magna. Those two species were chosen to assess whether the cyanobacteria-daphnid dynamics could be generalized for the Daphnia genus. Results demonstrated that both slope and EC50 of the concentration response curves depend upon the Daphnia species, the cyanobacteria species and the potential interaction between the two. This has two major consequences. First, the differences in sensitivity to cyanobacteria between D. magna and D. pulex depend upon concentration of the specific cyanobacteria. Second, we noted different mechanisms of toxicity for the two zooplankton species, a more general mechanism of toxicity for D. pulex and a more specific one for D. magna. Our data therefore suggest that results of studies investigating effects of cyanobacteria at different concentrations cannot be generalized across species. Furthermore, mechanisms of toxicity are not only cyanobacteria specific, but also dependent on the exposed species, even for rather closely related species such as in the Daphnia genus. Whenever possible, we therefore propose to combine a multi-species approach together with a full concentration response analysis to reach more general conclusions concerning the effects of cyanobacteria on zooplankton.     

Application of individual growth and population models of Daphnia pulex to Daphnia magna, Daphnia galeata and Bosmina longirostris

Daphnia models for individual growth and population dynamics have been developed in the manner of models developed by Gurney, McCauley, Andersen and others. All or most of the earlier models were parameterized for Daphnia pulex; we have used the D. pulex model as a baseline model for other species of Daphnia such as magna, galeata and also Bosmina longirostris. Because of the lack of ample data for D. magna, D. galeata and B. longirostris, some of the physiological data had to be relied on the other species whose data were available and in some case calibrated. We were able to produce reasonable results for individual growth as well as population dynamics under the controlled laboratory conditions. Most of the results were compared with the available laboratory data for population as well as growth. All the simulations have been done under high and low food concentrations. The animals are assumed to be feeding on green algae (Chlamydomonas reinhardtti) under the laboratory conditions of 18–20°C. The continuous growth until the end of the life was observed in smaller B. longirostris, whereas rapid growth in the beginning and slower after the start of the reproduction was observed in Daphnia species. The smaller species matured earlier than larger species. B. longirostris population sustained better than Daphnia species in medium food concentrations.     

Evolutionary history of contagious asexuality in Daphnia pulex

Asexual taxa are short-lived, suggesting that transitions to asexuality represent evolutionary dead-ends. However, with high rates of clonal origin and coexistence of asexuals and sexuals via selective asymmetries, asexuality may persist in the long term as a result of a dynamic equilibrium between clonal origin and extinction. Few such systems have been studied in detail. Here, we investigate the evolutionary history of asexual lineages of Daphnia pulex, which are derived from sexual relatives via the inheritance of a dominant female-limited meiosis-suppressing locus and inhabit ponds throughout northeastern North America (NA). Our extensive sampling and subsequent phylogenetic analysis using mitochondrial sequence data reveals a young and genetically diverse asexual assemblage, reflecting high rates of clonal origin due to the contagious nature of asexuality. Yet, asexuality is restricted to two phylogroups (B and C) with historical and/or present associations with northeastern NA and is absent from a northwestern phylogroup (A), supporting a recent northeastern origin of asexuality in this species. Furthermore, macrogeographic patterns of genetic variability indicate that phylogroups B and C recolonized northeastern NA from opposite directions, yet their presently overlapping geographic distributions are similarly divided into an eastern asexual and a western sexual region. We attribute these patterns to a recent contagious spread of asexuality from a northeastern source. If environment-mediated selective asymmetries play no significant role in determining the outcome of competitive interactions between sexuals and asexuals, regions of contact may be setting the stage for continued asexual conquests.     

Reproductive costs of Chaoborus-induced polymorphism in Daphnia pulex

Although the Chaoborus-induced spined morph of Daphnia pulex survives attacks by Chaoborus over twice as frequently as the typical morph, the spined morph is never found in the absence of Chaoborus. This implies that a disadvantage is associated with the spined morph in the absence of Chaoborus predation. The present study tested the hypothesis that the typical morph has a higher intrinsic rate of increase than the spined morph, by measuring several life history characteristics in controlled laboratory experiments at constant temperature and unlimited food.The results suggest that the spined morph of D. pulex takes longer to reach maturity, is smaller at maturity, but has similar egg number and egg size as the typical morph. These results are consistent with the hypothesis that the Chaoborus-induced spined morph is reproductively inferior to the typical morph.     

Demographic costs of Chaoborus-induced phenotypic plasticity in Daphnia pulex

Summary It has been proposed that morphological defenses against predation have demographic costs. We measured the cost of a predator-induced morphological defense, using predaceous phantom midge larvae Chaoborus americanus (Insecta, Diptera) and the prey species Daphnia pulex (Crustacea, Cladocera). The induced defense is a neck tooth (and other pleiotropic structures) developed in juvenile D. pulex in the presence of C. americanus. Laboratory life table experiments, in the absence of predation, indicated the population growth rate of typical D. pulex was 11% to 39% greater than that of D. pulex exposed to C. americanus extract, or C. americanus-conditioned water. The reduction in population growth rate was most frequently associated with an increase in the time between birth and first reproduction. Induced individuals required twenty more hours at 23°C, and twenty five more hours at 20°C, to develop to the age of first reproduction. Under limiting food conditions age-specific survivorship and the number of offspring produced per female by the induced form were reduced relative to the typical form. As a result, the difference in population growth rates among forms was greater at the low food level as indicated by a highly significant food by form interaction effect. In addition to neck teeth and lowered reproductive rates, the offspring of induced form individuals had significantly longer tail-spines (7.2–7.5%), and primiparous adults from the induction treatment were significantly shorter than controls (3–8%).     

Sexual reproduction of Daphnia pulex in a temporary habitat

Species of Daphnia (Crustacea: Cladocera) typically reproduce by cyclical parthenogenesis, in which a period of all-female parthenogenetic reproduction is followed by sexual reproduction. Sex in Daphnia is determined by the environment, with factors such as temperature, photoperiod and crowding stimulating the production of males and sexual females. Previous studies on Daphnia pulex from temporary pond habitats demonstrated the coexistence of male-producing and non-male-producing (NMP) females, as determined under crowding in the laboratory. A strong genetic component to this sex allocation variation suggested that sex expression in D. pulex is better described as a result of genotype-environment interaction. The present study examined the switch from parthenogenetic to sexual reproduction in two temporary-pond populations of D. pulex. Both populations showed a very early investment in sexual reproduction, independent of population density, by producing males very soon after the populations were reestablished from resting eggs in the early spring. Approximately 40% of the initial broods were male. Additional evidence for gender specialization was obtained by observing the sex of two or three successive broods for 85 individual females. Fifty-eight females produced successive broods of females, 13 females produced successive broods of males and 14 females produced successive broods which included both male and female broods. Females that produced successive female broods under natural conditions included a higher frequency of NMP females compared to a random sample of females, confirming the existence of NMP females. Sexual females were observed in both populations after the first appearence of males, suggesting that the presence of males may stimulate the production of sexual females. For D. pulex populations in a temporary environment, there appears to be an increased emphasis on sexual reproduction and a decreased influence of the environment on sex determination, compared to Daphnia populations in more permanent habitats. Received: 19 February 1996 / Accepted: 20 January 1997