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.     

Ecological differences among clones of Daphnia pulex Leydig

Summary Natural populations of Daphnia pulex that reproduce by obligate parthenogenesis include a number of clones. Studies on two common and two rare clones from southwestern Ontario revealed significant differences in their intrinsic rates of increase, competitive abilities, rates of ephippial egg production, and lifespans. Environmental factors such as temperature and food type had large inluences on the rate of increase of each clone. Differences in rates of increase among clones were most pronounced at temperatures higher than those encountered in nature. In general, the covariance of life history traits among clones was high. The outcome of competitive encounters between clones was deterministic and in most cases was unaffected by temperature. Clones with high rates of increase tended to be better competitors than those with low rates of increase.     

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.     

Significance of photoperiodism and diapause control in the multicycle Crustacean Daphnia pulex Leydig

Eukaryote cells traverse the cell cycle, or become growth-arrested at one of two blockpoints, as in an engine of energy conversion. Diapause describes that growth arrest in the cell cycle(s), of single and multicellular individuals. Its temporal position in an annual cycle is an evolved response to the external environment, which is also responsible for sending the necessary signal(s) to receptors in the cell, or to the neural integrator of multicellular individuals.The diapause syndrome, common to protists and Crustacean waterfleas, includes the cell cycle arrest, as well as the induction of mating types, gametogenesis, packaging of the diapaused stage, and the duration of the diapause. The cell-/ life-cycle may be interrupted in a variety of seasonal patterns, which are triggered by one or more identified external signals and are integrated by cell and circadian clocks. Daphnia pulex displays four distinct seasonal patterns of embryonic diapause in four putative genotypes.The ecological implications of diapause in life cycles may be sought in such questions as 1, environmental control of temporal and spatial boundaries in sessile populations; 2, application of knowledge of diapause/dormancy cycles to redesigning ecological communities; and 3, integration of mechanistic and biosystematic approaches in ecology.     

Competitive abilities and distribution ranges of four asexual Daphnia pulex lineages and Daphnia mitsukuri in Eurasian continental islands

Daphnia pulex and D. mitsukuri are morphologically similar and distributed in small lowland Japanese lakes. Daphnia pulex in Japan reproduces by obligate parthenogenesis and is composed of four different lineages (JPN1–JPN4) with North American origins. Although this species is found throughout Japan, JPN1 has the largest distribution range, followed by JPN2. Daphnia mitsukuri is a putatively endemic species that is now rarely found in Japan. Since Daphnia share the same algal food, the difference in distribution ranges among the four asexual D. pulex lineages and D. mitsukuri may be caused by exploitative competition for algal food. To examine this possibility, we measured the threshold food concentration (TFC) at zero-net growth rate and the food concentration required for 50% of the individuals to survive (FC50). In addition, we examined the effects of temperature on the somatic growth rate (SGR) and the mortality rates of four asexual D. pulex lineages and D. mitsukuri. The experiments showed that TFC was lowest in D. pulex JPN1 and highest in D. mitsukuri, supporting the idea that the largest distribution range of JPN1 is due to its superiority in inter-lineage competition, and that the distribution area of D. mitsukuri is reduced through exploitative competition with D. pulex. However, although JPN2 was found in relatively large areas, it had higher TFC than JPN3 and JPN4, which appear in very limited areas. These results clearly showed that the difference in distribution ranges among D. pulex lineages are not necessarily determined by their competitive superiority alone.     

Variation in allocation to sexual and asexual reproduction among clones of cyclically parthenogenetic Daphnia pulex (Crustacea: Gladocera)

Organisms reproducing by cyclical parthenogenesis combine the benefits of both sexual and asexual reproduction within the same life cycle. Few studies have examined the evolution of variation in the pattern of investment in parthenogenetic compared to sexual reproduction. Seven clones of Daphnia pulex (Crustacea: Cladocera) varying in allocation to sexual reproduction, as measured by the production of males, were raised in isolation and together in a microcosm to study the pattern of sexual reproduction and the effect of this variation on clone fitness. Sex allocation for clones raised together a microcosm was similar to their allocation when raised in isolation, suggesting a genetic basis to the variation. Three clones showed a cost of producing males that lead to their extinction after about 30 days due to the lack of females required for the clones to persist by parthenogenetic reproduction. The remaining four clones persisted until the end of the 72-day experiment. Clones with little or no allocation to males showed no increased allocation to sexual females. The seven clones showed a greater variation in estimated fitness through male and female function than in total estimated fitness. The clone with the greatest total fitness gained most of its fitness through male function but also had a relatively high fitness through female function. Although one clone produced only females it had the next highest fitness. The three clones that went extinct because of a high investment in males had estimated fitness as high as some clones that persisted in the microcosm because of a higher investment in parthenogenetic reproduction. The similarity in total fitness among clones suggests that Daphnia pulex populations in temporary habitats maintain a sex polymorphism where different genotypes vary-in functional gender ranging from female to primarily male.     

Life history variation among low-arctic clones of obligately parthenogenetic Daphnia pulex: a diploid-polyploid complex

Summary Laboratory life table experiments were conducted using nine clones of obligately parthenogenetic Daphnia pulex that were collected from a site in the Canadian low-arctic. Two of the nine clones were diploids, while the other seven clones were polyploids. Significant clonal differences in age at first reproduction, size at first reproduction, number of offspring in each of the first three broods, offsrring sizes for the first two broods, and intrinsic rates of natural increase were detected. Differences in life histories were evident between polyploids and diploids. Generally, polyploid clones reached maturity at later ages, matured at larger sizes, produced smaller broods, and larger offspring than the diploid clones. The data are discussed in reference to potential biotic (i.e. invertebrate predation) and abiotic factors (i.e. physicochemical gradients) that may influence life history variation in this clonal assemblage.     

Ammonia-nitrogen excretion in Daphnia pulex

Ammonia-nitrogen excretion rates were measured in natural summer and cultured populations of Daphnia pulex from Silver Lake, Clay County, Minnesota, USA during 1973. The mean rate of ammonia-nitrogen excretion for the summer populations was 0.20 µg N animal^–1 day^–1 or 5.11 µg N mg^–1 dry body weight day^–1 (N = 80) measured at 15°, 20°, and 25°C. These rates appear to be temperature and weight dependent, but they are probably affected by factors other than temperature and dry body weight. Ammonia-nitrogen excretion rates of Daphnia pulex cultured on Chlamydomonas reinhardi yielded the following relationship with temperature: Log₁₀E = (0.061) T 1.773, where E is µg N animal^–1 day^–1 and T is temperature °C. The ammonia-nitrogen excretion on a mg^–1 dry body weight day^–1 basis was related to temperature according to the following similar expression Log₁₀E = (0.043) T + 0.153, where E is µg N mg^–1 dry body weight day^–1, and T is temperature °C. The length-weight relationship of Daphnia pulex for the summer populations (N = 1583) was log₁₀W = (0.526) Log₁₀L + 1.357, where W is weight in µg and L is length in mm.     

Eye movements in Daphnia pulex (De Geer).

1. The various types of eye movement exhibited by the cyclopean eye of Daphnia pulex were studied using high speed motion photography. 2. This rudimentary eye, which consists of only 22 ommatidia, can move through approximately 150 degrees in the sagittal plane and 60 degrees in the horizontal plane. 3. Four classes of eye movement were found: (1) a high speed tremor at 16 Hz with an amplitude of 3-4 degrees, which resembles physiological nystagmus, (2) a slow rhythmic scanning movement at 4 Hz, and 5-6 degrees amplitude, (3) large fast eye movements similar to saccadic eye movements and (4) optokinetic nystagmus produced by moving striped patterns. 4. Where the fast tremor occurred concurrently with the slow rhythmic scan, a Fourier analysis revealed that the former was the fourth harmonic of the latter.     

Sex allocation variation in Daphnia pulex

Daphnia (Crustacea: Cladocera) reproduce by cyclical parthenogenesis in which the sex of offspring is environmentally determined. Although numerous studies have demonstrated that factors such as crowding and short-day photoperiod stimulate male production, there is limited information on variation in allocation to male and female offspring for any species of Daphnia . The present study assessed the presence or absence of male production in 96 isofemale lines (clones) from each of eight populations of Daphnia pulex . An average of 37% (range 18–51%) of clones failed to produce males under crowded conditions in the laboratory. A subset of 14 of these non-male-producing clones also failed to produce males under short-day photoperiod (8L:16D). Three male-producing clones were within-clone mated as well as crossed to three non-male-producing clones to study the inheritance of the failure to produce males. The average frequency of non-male-producing F ₁ progeny was significantly higher (58%, N = 486) among the outcrossed progeny than the inbred progeny (5%, N = 86). In addition, when sixteen of the male-producing outcrossed progeny were within-clone mated, only 7% ( N = 106) of the resulting F ₂ progeny failed to produce males. These results are consistent with a genetic basis for the absence of male production. Average survival of the progeny from the nine outcrossed matings was more than twice (67%) that of the inbred progeny from the three within-clone matings (30%), suggesting that within-clone mating would result in significant inbreeding depression. We present a model that suggests that even low levels of inbreeding could allow non-male-producing females to be maintained in a population. The co-occurrence of non-male-producing females and females that produce both males and females in Daphnia pulex bears a similarity to the gynodioecious breeding system found in some plant species.     

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.     

Size-specific swimming behavior of Daphnia pulex

Body length affects several aspects of the behavior of quietlyswimming Daphnia pulex Swimming and sinking rates were measuredat 0.033 s intervals during the ‘hops’ characteristicof Daphnia swimming behavior Larger animals swim faster, covermore distance, and produce more powerful swimming strokes. LargerDaphnia also sink faster, but the sinking rate scales as lengthto the 0 58 power, far lower than the power of 2 00 predictedby Stokes Law considerations. The number of hops s^–1 wasindependent of body size, although a theoretical analysis predictshopping rate (antennal beat frequency) should increase as bodylength squared. Turning behavior, measured as the ratio of displacementto total distance, during 5 s, is also independent of body sizeIndependence of several parameters of body motion and body sizeimplies that factors other than simple mechanics affect Daphniaswimming behavior     

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.     

Genome size evolution and polyploidy in the Daphnia pulex complex (Cladocera: Daphniidae)

Genome size was estimated in 49 clones of the Daphnia pulex complex from temperate and subarctic locations using flow cytometry and microsatellite DNA analyses. Significant genome size differences were found in diploid species belonging to the two genetically distinct groups (the pulicaria and the tenebrosa groups), with clones from the tenebrosa group having genome sizes 22% larger than those in the pulicaria group. Combined flow cytometry and microsatellite DNA analyses revealed that nearly all polyploid clones in the D. pulex complex are triploid and not tetraploid, as was previously suggested. Sequencing analyses of the ND5 gene to position clones in their respective clades within the D. pulex complex have uncovered three triploid clones of Daphnia middendorffiana with a D. pulex maternal parent. This result was unexpected because Daphnia pulicaria has always been identified as the maternal parent of these hybrid polyploid clones. Triploid clones likely owe their origins to interactions between sexual and asexual populations. Further interactions in the tenebrosa group have generated tetraploid clones but these events have been rare.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 68–79.     

Feeding rate inhibition in crowded Daphnia pulex

Feeding rates of Daphnia pulex fed a range of levels of the alga Chlamydomonas reinhardi of 15 °C are strongly density-dependent. At lower densities, Daphnia (30 1^–1) fed at higher rates than crowded (270 1^–1) Daphnia which manifest a relatively depressed saturation feeding response. At 30 individuals/liter, Daphnia consumed 8.5 – 15.7 × 10⁴ cells d^–1h^–1 (on a volume basis, 12.1 – 22.2 × 10⁶ m³), at 270 L^–1 3.7 – 3.9 × 10⁴ (5.2 – 5.5 = 10⁶ m³ cells d^–1h^–1 when feeding on algae at 80 000 cells ml^–1 (11.3 × 10⁶ m³ ml^–1). The feeding rate data best fit an Ivlev feeding function. An autoallelopath might be causing the repression. Water preconditioned with crowded Daphnia completely repressed feeding in uncrowded Daphnia after six hours.     

Activation of the Ephippial Egg of Daphnia pulex

The ephippial eggs of Daphnia pulex require light for the initiation of development. The ephippial capsule prevents the completion of development but is not a barrier to an adequate light stimulus. Working with decapsulated eggs, the response to light increased to 100 % within 9 days of storage in the dark and remained at 100% for up to 60 days of storage in the dark. The response was not dependent on drying the ephippia. Ephippia stored in the light did not reach 100% response to illumination when decapsulated, indicating that activation was dependent on prior dark reactions. About 4500 ft-c-min of fluorescent light energy was required for 100% activation. The effective wavelengths were between 350 and 475 mµ with 2 x 10⁶ ergs/cm² sufficient to initiate nearly 100% development at 410 mµ, the most effective wavelength. Low temperature interfered with photoactivation but not with subsequent development. Chilling the ephippia resulted in an increased light requirement. Kinetic studies with chilled ephippia stored for various times in the dark indicated a diphasic process of photoactivation which has tentatively been interpreted as a light-dependent release of inhibition.     

Transcontinental Phylogeography of the Daphnia pulex Species Complex

Daphnia pulex is quickly becoming an attractive model species in the field of ecological genomics due to the recent release of its complete genome sequence, a wide variety of new genomic resources, and a rich history of ecological data. Sequences of the mitochondrial NADH dehydrogenase subunit 5 and cytochrome c oxidase subunit 1 genes were used to assess the global phylogeography of this species, and to further elucidate its phylogenetic relationship to other members of the Daphnia pulex species complex. Using both newly acquired and previously published data, we analyzed 398 individuals from collections spanning five continents. Eleven strongly supported lineages were found within the D. pulex complex, and one lineage in particular, panarctic D. pulex, has very little phylogeographical structure and a near worldwide distribution. Mismatch distribution, haplotype network, and population genetic analyses are compatible with a North American origin for this lineage and subsequent spatial expansion in the Late Pleistocene. In addition, our analyses suggest that dispersal between North and South America of this and other species in the D. pulex complex has occurred multiple times, and is predominantly from north to south. Our results provide additional support for the evolutionary relationships of the eleven main mitochondrial lineages of the D. pulex complex. We found that the well-studied panarctic D. pulex is present on every continent except Australia and Antarctica. Despite being geographically very widespread, there is a lack of strong regionalism in the mitochondrial genomes of panarctic D. pulex – a pattern that differs from that of most studied cladocerans. Moreover, our analyses suggest recent expansion of the panarctic D. pulex lineage, with some continents sharing haplotypes. The hypothesis that hybrid asexuality has contributed to the recent and unusual geographic success of the panarctic D. pulex lineage warrants further study.