Colonization dynamics and life history traits of seven Daphnia pulex genotypes

Summary A series of experiments revealed significant differences in the potential ability of seven Daphnia pulex genotypes to colonize two lakes in which this species does not naturally reside. Life table experiments, in which individuals from each genotype were raised separately on water and natural phytoplankton from the two lakes, revealed several significant differences among genotypes in life history traits, including age and size at first reproduction, clutch size and offspring body size. Significant differences among genotypes were also found in mean genotype fitness and rate of population increase, although all genotypes were able to increase in absolute numbers. Significant genotypexlake interaction was found for several life history traits and mean fitness, indicating that the relative success of invading genotypes may depend on habitat characteristics. Enclosure experiments, in which all seven genotypes were introduced together into enclosures in both lakes, revealed that some genotypes increased greatly while others declined in relative abundance, and that the most successful genotypes differed between lakes. In addition, the most successful genotypes in the enclosures were not necessarily the genotypes that displayed the highest fitness in the life table experiments, possibly because individuals in the enclosures competed for food resources, leading to exclusion of certain genotypes.     

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     

Microsporidiosis in a Daphnia pulex population

The impact of parasitism by Thelohania sp., an intracellular microsporidian, on Daphnia pulex inhabiting a vernal pond was studied for three years. Three issues were considered: the distribution of parasites among hosts, the effect of parasites on individuals, and the impact of parasitism on population growth. Each year, Thelohania infected members of at least half of the generations of Daphnia produced in the pond. When the frequency of infection was low, parasites were found only in large adults. As infection frequency increased, parasitism spread downward through host size classes. However, parasites were rarely found in juveniles. Parasitism reduced clutch size drastically, increased mortality to a variable extent, had little impact on egg size or on per instar growth and none on molt frequency. Interaction with other stresses, such as food limitation, exacerbated some of these effects. Parasitism reduced instantaneous birth rate much more than it elevated instantaneous death rate. Population growth was reduced significantly but it is unlikely that Thelohania alone regulates the growth of this Daphnia population.     

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.     

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.     

Temperature and kairomone induced life history plasticity in coexisting Daphnia

We investigated the life history alterations of coexisting Daphnia species responding to environmental temperature and predator cues. In a laboratory experiment, we measured Daphnia life history plasticity under different predation risk and temperature treatments that simulate changing environmental conditions. Daphnia pulicaria abundance and size at first reproduction (SFR) declined, while ephippia (resting egg) formation increased at high temperatures. Daphnia mendotae abundance and clutch size increased with predation risk at high temperatures, but produced few ephippia. Thus, each species exhibited phenotypic plasticity, but responded in sharply different ways to the same environmental cues. In Glen Elder reservoir, Kansas USA, D. pulicaria dominance shifted to D. mendotae dominance as temperature and predation risk increased from March to June in both 1999 and 2000. Field estimates of life history shifts mirrored the laboratory experiment results, suggesting that similar phenotypic responses to seasonal cues contribute to seasonal Daphnia population trends. These results illustrate species-specific differences in life history plasticity among coexisting zooplankton taxa.     

Seasonal variation of egg size and number in a Daphnia pulex population

Seasonal variation of egg size and number was examined in a Daphnia pulex population inhabiting a vernal pond. In this population, size at maturity declines at midseason, probably as an adaptive response to size-selective predation by larvae of the salamander Ambystoma. The larger early season individuals produce more and larger eggs than the smaller late season individuals. Age at maturity does not vary between seasons. Laboratory experiments indicate that temperature may affect egg size, egg number and size at maturity. However, field data suggest that temperature accounts for only a small fraction of the total variation in egg size and number. Indirect measures of nutrition indicate that food limitation does not cause the seasonal decline in egg size and number. The seasonal change in reproductive traits is well correlated with changes in invertebrate and vertebrate predation. Examination of predator feeding preferences and their impact on Daphnia mortality indicate that variation of reproductive traits is most likely a complex adaptation to changing predation regimes.     

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.     

Progress towards characterization of a predator/prey kairomone: Daphnia pulex and Chaoborus americanus

Predatory larvae of the midge Chaoborus americanus induce an antipredator defense (neckteeth) in prey individuals of the cladoceran Daphnia pulex. The signal for presence of predator is a water-soluble chemical. We provide evidence that this kairomone originates in the intestinal tract of the predator. The active compound is an organic molecule of intermediate polarity which is heat stable and partially destroyed by acid and base digestion. It is stable to digestion by the general peptidase Pronase. Hydroxyl groups, but not primary amines, carbonyls or thiols are essential to activity. Low-pressure liquid chromatography on a reverse-phase silica gel (Amicon Matrex C-18) column suggests there may be more than one active component.     

Esterase genes in Daphnia pulex: linked inheritance and genotypic distribution in natural populations

Summary Esterase patterns were examined in three populations of Daphnia pulex. The total number of bands showing esterase activity was 17. Three major genes Est-1, Est-2, Est-3 controlling esterase synthesis were identified and genetically studied. These genes were found to be located in the same linkage group. It was shown that two or three homologous chromosomes differing in sets of esterase alleles predominantly occur in the populations considered.     

Effects of Microcystis aeruginosa on interference competition between Daphnia pulex and Keratella cochlearis

Laboratory experiments tested the hypothesis that a toxic strain of Microcystis aeruginosa decreases the ability of Daphnia pulex to interfere with Keratella cochlearis. To test a variety of conditions, juvenile and adult Daphnia were exposed to the cyanobacterium for different time periods prior to, and during the experiments. Adult Daphnia not only suppressed rotifers over successive two-day intervals, but also had a significant impact within a 24-hour period. However, the presence of Microcystis (5 × 105 cells ml^–1) decreased the Daphnia effect in both experiments. Although juvenile Daphnia also significantly suppressed Keratella population growth, the presence of Microcystis (10⁵ and 5 × 10⁵ cells ml^–1) caused a significant reduction in daphniid body size and decreased the ability of both nonacclimated and acclimated daphniids to suppress rotifers. Keratella inhalation and mortality are positively correlated with filtering rates and body size of Daphnia. Therefore, the feeding rates and size structure of a Daphnia population will determine its potential to interfere with vulnerable rotifers. In all experiments the presence of Microcystis significantly decreased the ability of Daphnia to interfere with this rotifer despite the fact that Keratella was also inhibited. In the field this effect might be augmented if Microcystis colonies are more easily ingested by cladocerans than by the rotifers.     

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.     

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.     

Effects of predator-released chemicals on some life history parameters of Daphnia pulex

The effects of chemicals released by fish and Chaoborus larvae on some life history traits in Daphnia pulex were studied in an in situ enclosure experiment. The mean size of Daphnia individuals was smaller in the presence of fish-released cues. Also the minimal size of an egg bearing female in the presence of fish exudates was smaller than in the population exposed to the chemicals released by Chaoborus larvae as well as in the control population. Fish-released chemicals caused the increase in clutch size in Daphnia. There were no statistically significant differences between the studied life history parameters of the control and Chaoborus treatments. The results are discussed in reference to recent laboratory research.     

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.     

Food quality effects on life history traits and fitness in the generalist herbivore Daphnia

Summary Life table experiments were conducted on the generalist suspension feeder Daphnia galeata, using as food the two green algae (Chlorophyta) Scenedesmus acutus and Oocystis lacustris. Oocystis was hypothesized to be a lower quality food because it is convered with a thick sheath, believed to reduce digestibility. Results showed that Oocystis is a lower quality food for Daphnia, but only at relatively low food concentrations (0.15 mg C/L) and not at higher concentrations (1.0 mg C/L). At 0.15 mg C/L, Daphnia intrinsic rate of increase (r) when grown on Oocystis was only half that when grown on Scenedesmus. Daphnia r was similar at 0.15 mg C/L Oocystis and 0.075 mg C/L Scenedesmus, indicating that Daphnia requires twice as much Oocystis as Scenedesmus to achieve the same fitness. Intrinsic rate of increase was lower on Oocystis mainly because age at first reproduction was greatly delayed compared to that on Scenedesmus (13.6 vs 7.3 d). In addition, juvenile growth and survivorship were reduced on Oocystis compared with Scenedesmus. Clutch sizes were similar on the two foods, indicating that once individuals reached adulthood, the two foods were similar in quality. In contrast, at high food concentrations (1.0 mg C/L), the two algae were similar in quality for both juveniles and adults, and r was not significantly different on the two foods. Ingestion and assimilation rate experiments whowed that Daphnia consumes the two algae at identical rates, and that adults assimilate the two algae at similar rates. However, juveniles assimilate Oocystis at much lower rates than Scenedesmus, possibly accounting for reduced juvenile growth and delay in age at maturity at low concentrations. Thus, Daphnia exhibits an ontogenetic shift in its ability to utilize Oocystis, and this can result in juvenile bottlenecks in which survival and growth of young age classes are of critical importance in determining population dynamics. Because food quality effects were manifested primarily in juveniles and at low concentrations, food quality effects in nature will depend on phytoplankton abundance and age-structure of Daphnia populations.     

A population of sexual Daphnia pulex resists invasion by asexual clones

Asexual reproduction avoids the costs associated with sex, predicting that invading asexual clones can quickly replace sexual populations. Daphnia pulex populations in the Great Lakes area are predominately asexual, but the elimination of sexual populations by invading clones is poorly understood. Asexual clones were detected at low frequency in one rare sexual population in 1995, with some increase in frequency during 2003 and 2004. However, these clones remained at low frequency during further yearly sampling (2005–2013) with no evidence that the resident sexual population was in danger of elimination. There was evidence for hybridization between rare males produced by asexual clones and sexual females with the potential to produce new asexual genotypes and spread the genetic factors for asexuality. In a short-term laboratory competition experiment, the two most common asexual clones did not increase in frequency relative to a genetically diverse sexual population due in part to a greater investment in diapausing eggs that trades-off current population growth for increased contribution to the egg bank. Our results suggest that a successful invasion can be prolonged, requiring a combination of clonal genotypes with high fitness, persistence of clones in the egg bank and negative factors affecting the sexual population such as inbreeding depression resulting from population bottlenecks.     

The effects of Chilomonas on the life history traits of [-2pt] Daphnia longispina under semi-natural conditions and the implications for competition in the plankton

Assuming that heterotrophic flagellates (H.F.) can sustain cladoceran life cycles, particularly at periods of low food and high detritus conditions, the growth, reproduction and life span of Daphnia longispinawere studied under conditions of summer food limitation. They were fed both natural resources and natural food enriched with a culture of the colourless Chilomonas (ovoid cell, 8 × 25 m). Four H.F. morphotypes occurred in the natural water and the first experiments with Daphnia, showed that the cladoceran would most easily ingest those of 5 m–10 m, while the addition of Chilomonas severely depressed the H.F. of 2 m. The capability of this flagellate to ingest small H.F. was confirmed using fluorescent particles ranging from 0.94 to 3.95 m. As a consequence, Daphnia could control the abundance of H.F., but also compete with the largest morphotypes. To study the influence of the H.F. on the life span of Daphnia, the cladoceran was acclimated for two generations, before the start of the experiments. In semi-natural conditions, constant temperature, without predators and fed natural water, the summer daphnids achieved smaller sizes, produced fewer offspring and lived for a shorter time than when Chilomonas was added to their diet. Despite the addition of 560 g C l^-1, this was not sufficient to enhance the Daphnia reproduction to the level of the spring population fed natural water. These results clearly illustrate that the heterotrophic flagellates contributed significantly to improving the reproduction and survival of the daphnids. They emphasise the importance of the microbial loop as a link to larger consumers, particularly when unpalatable or nutritionally inadequate algae are dominant, which is often so in lakes during summer.     

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.