Genetic versus environmental influence on pigment variation in the ephippia of Daphnia pulicaria

1. The dormant eggs of Daphnia (Crustacea: cladocera) are encased in a protective, chitonous casing known as an ephippium. Ephippia are pigmented with melanin, and the degree of pigmentation ranges from transparent to opaque. Variation in pigmentation exists within and across populations and species, raising questions about the factors that influence the natural distribution of pigmentation. 2. We used image analysis to quantify pigmentation in ephippial casings of Daphnia pulicaria that were produced both in the field and in the laboratory. The percentage of the surface area of ephippia that is darkly pigmented ranged from 0.5 to 99.5%. The range, mean and variance of ephippial pigmentation varied across our 11 study lakes. 3. Laboratory experiments compared clonal variation (five clones/lake) and population variation (five lakes) at three temperatures (15 °C, 20 °C and 25 °C). The degree of pigment variation between clones was much greater than pigment variation between temperatures, indicating that there is a genetic component to pigmentation in the ephippia which is stronger than the effect of temperature. 4. Comparisons of pigment levels and lake characteristics were used to identify physical and biological factors associated with ephippial pigmentation. Investment in ephippial production was the strongest predictor with darker ephippia occurring more often in lakes with the highest ephippial production.     

Molecular genetic analysis of Daphnia in the English Lake District: species identity, hybridisation and resting egg banks

• 1 Taxonomic issues in many Daphnia species complexes are often confused by a high degree of phenotypic plasticity and by interspecific hybridisation. Here, we employ molecular genetic tools to confirm the species composition and incidence of hybridisation in extant and resting egg populations of Daphnia from Windermere and Esthwaite Water in the English Lake District.
• 2 A combination of species‐diagnostic allozymes and mtDNA, confirms that contemporary populations are dominated by a single species, D. galeata.
• 3 We present the first account of genetic characterisation of dated ‘resting’ eggs using microsatellites and mtDNA, employing PCR‐based DNA recovery, thus providing a temporal dimension to taxonomic patterns. Thirty years ago, two species were present in Esthwaite, D. galeata and D. hyalina, but Windermere populations were dominated by D. galeata only.
• 4 The use of PCR‐based mtDNA RFLP analysis as a species‐diagnostic tool, and microsatellites to monitor clonal diversity, provide a valuable approach for long‐term studies, especially in populations free from the complicating effect of frequent hybridisation. The detailed limnological records available for many large lakes, and associated changes in land‐use, pollutants and climate, combined with long‐term ephippial molecular genetic data, provide opportunities for exploring natural and anthropogenic impacts on genetic and community structure.

     

How well do laboratory experiments explain field patterns of zooplankton emergence?

1. We conducted a laboratory experiment to explore potential mechanisms driving variation in zooplankton emergence from diapausing eggs observed in Oneida Lake, NY, U.S.A. We hypothesized that variation in timing of ice-out (date of thawing of ice) between 1994 and 1995, which resulted in variation in photoperiod–temperature cues, contributed to the differences in the observed field patterns. Environmental chambers were used to establish weekly photoperiod–temperature combinations that reproduced natural conditions in Oneida Lake in 1994 and 1995. In addition, a third treatment ('dark') exposed eggs only to the changes in temperature. We recorded zooplankton emergence for 2.5 simulated ice-free seasons.
2. Nine cladoceran taxa were found to hatch, but only Daphnia pulicaria in large numbers. Hatching of D. pulicaria was recorded throughout the season in the two light treatments and sporadically in the dark treatment. The early ice-out treatment had the highest emergence, followed by the late ice-out and dark treatments. Among taxa, there was temporal segregation with five hatching in the early weeks of sampling and two taxa hatching during the middle weeks. Alona hatched late in the first year, but earlier in the second year.
3. We compared our laboratory results of D. pulicaria hatching with the field data obtained by Cáceres (1998) . Hatching was continuous in the laboratory, whereas a synchronous spring emergence was found in the field. However, in both the laboratory and the field, more D. pulicaria hatched under conditions reflecting ice-out occurring in March as opposed to April. Because differences in rates and timing of emergence can affect the population and community dynamics of pelagic systems, we suggest caution when applying laboratory results to field populations.     

Predator-induced phenotypic plasticity in the exotic cladoceran Daphnia lumholtzi

1. The exotic cladoceran Daphnia lumholtzi has recently invaded freshwater systems throughout the United States. Daphnia lumholtzi possesses extravagant head spines that are longer than those found on any other North American Daphnia. These spines are effective at reducing predation from many of the predators that are native to newly invaded habitats; however, they are plastic both in nature and in laboratory cultures. The purpose of this experiment was to better understand what environmental cues induce and maintain these effective predator‐deterrent spines. We conducted life‐table experiments on individual D. lumholtzi grown in water conditioned with an invertebrate insect predator, Chaoborus punctipennis, and water conditioned with a vertebrate fish predator, Lepomis macrochirus. 2. Daphnia lumholtzi exhibited morphological plasticity in response to kairomones released by both predators. However, direct exposure to predator kairomones during postembryonic development did not induce long spines in D. lumholtzi. In contrast, neonates produced from individuals exposed to Lepomis kairomones had significantly longer head and tail spines than neonates produced from control and Chaoborus individuals. These results suggest that there may be a maternal, or pre‐embryonic, effect of kairomone exposure on spine development in D. lumholtzi. 3. Independent of these morphological shifts, D. lumholtzi also exhibited plasticity in life history characteristics in response to predator kairomones. For example, D. lumholtzi exhibited delayed reproduction in response to Chaoborus kairomones, and significantly more individuals produced resting eggs, or ephippia, in the presence of Lepomis kairomones.     

Daphnia community analysis in shallow Kenyan lakes and ponds using dormant eggs in surface sediments

1. Water fleas of the genus Daphnia are considered rare in tropical regions, and information on species distribution and community ecology is scarce and anecdotal. This study presents the results of a survey of Daphnia species distribution and community composition in 40 standing waterbodies in southern Kenya. The study sites cover a wide range of tropical standing aquatic habitats, from small ephemeral pools to large permanent lakes between approximately 700 and 2800 m a.s.l. Our analysis combines data on Daphnia distribution and abundance from zooplankton samples and dormant eggs in surface sediments. 2. Nearly 70% (27 of 40) of the sampled waterbodies were inhabited by Daphnia. Although their abundance in the active community was often very low, this high incidence shows that Daphnia can be equally widespread in tropical regions as in temperate regions. 3. Analysis of local species assemblages from dormant eggs in surface sediments was more productive than snapshot sampling of zooplankton communities. Surface‐sediment samples yielded eight Daphnia species in total, and allowed the detection of Daphnia in 25 waterbodies; zooplankton samples revealed the presence of only four Daphnia species in 16 waterbodies. 4. Daphnia barbata, D. laevis, and D. pulex were the most frequently recorded and most abundant Daphnia species. Canonical correspondence analysis of species–environment relationships indicates that variation in the Daphnia community composition of Kenyan waters was best explained by fish presence, temperature, macrophyte cover and altitude. Daphnia barbata and D. pulex tended to co‐occur with each other and with fish. Both species tended to occur in relatively large (>10 ha) and deeper (>2 m) alkaline waters (pH 8.5). Daphnia laevis mainly occurred in cool and clear, macrophyte‐dominated lakes at high altitudes.     

The importance of hatching from resting eggs for population dynamics and genetic composition of Daphnia in a deep reservoir

1. Cyclical parthenogens, organisms switching between sexual and asexual mode of reproduction, are common in aquatic habitats. In permanent waterbodies with year‐round persistence of populations, the relevance of sexually produced resting eggs for population dynamics and structure is still largely unexplored. 2. The aim of this study was to investigate the importance of recruitment from resting eggs to Daphnia spp. in a deep dimictic reservoir with overwintering populations (Saidenbach Reservoir, Germany). We linked field studies on Daphnia population dynamics, hatching rates and diapausing egg abundances in the sediment with allozyme analysis of the genetic variability of both the pelagic population and hatchlings from resting eggs. 3. We found higher hatching rates at intermediate depths (10–18 m), although highest diapausing egg abundances were found in the deeper parts of the reservoir. From the extensive field data, we calculated a population budget for the main basin of the reservoir, which showed that hatched daphnids accounted for <1% of the total abundance and explained only a tiny fraction of population growth. 4. The genetic composition (MultiLocus Genotypes, MLG) of the pelagic population was relatively stable throughout the stratified period with up to five among 39 detected MLGs attaining frequencies >5% per date. From 11 MLGs differentiated among hatched daphnids, three were recovered in the pelagic population in perceptible, albeit low frequencies (1–4%). 5. We conclude that the contribution of hatched animals to the spring population dynamics is negligible in situations with an overwintering population present and will only become relevant if densities are extremely low. In this type of system, the significance of the resting egg bank is presumably mainly related to the maintenance of genetic diversity.     

Diapause in Monogonont Rotifers

This review focuses on more recent findings on the diapause in Monogonont rotifers, since the major reviews by Pourriot & Snell (1983, Hydrobiologia 104: 213–224) and Gilbert (1992, Rotifera. In Adiyodi, K. G. & R. G. Adiyodi (eds), Reproductive Biology of Invertebrates, Vol. 5 – Sexual Differentiation and Behaviour. IBH Publishing Co., Oxford: 115–136; Vol. 6A – Asexual Propagation and Reproductive Strategies. IBH Publishing Co., Oxford: 231–263.). It covers diapausing egg formation, diapausing egg survival, and diapausing egg hatching as well as possible strategies involved in these processes. Data from laboratory and field studies exist, but little information is available on diapausing egg hatching in the field. Resting or diapausing egg production can be mictic and in some cases amictic. Mictic diapausing egg production depends on the balance between cues promoting and inhibiting mictic female production. Such cues can be either environmental or endogenous. Our knowledge on factors inducing mixis is still limited to a few species, but effects of crowding on mixis induction may be more widespread. Recent results also show that male mating behavior may play an important role in the successful production of diapausing eggs. Hatching may be induced by factors related to temperature and light conditions; also desiccation may have a role. However, desiccation of temporary habitats may also be damaging to diapausing stages. Although few data are available, those existing point to distinct differences between populations and the importance of clonal variation within populations regarding the initiation as well as the termination of diapause.     

To sink or float: the fate of dormant offspring is determined by maternal behaviour in Daphnia

• 1 As the ephippia (chitinous shells enclosing diapausing eggs) of pelagic crustaceans of the genus Daphnia have been occasionally reported to float at the water surface, we considered that this might be an adaptation promoting their passive dispersal. We investigated the mechanisms by which ephippia appear at the water surface.
• 2 While field surveys revealed that floating Daphnia ephippia are often numerous in various freshwater habitats, laboratory tests showed that newly formed ephippia are not buoyant initially. Once transferred to the surface by whatever means, however, they may remain there due either to surface tension or gas absorption.
• 3 Video recordings showed that all ephippia at the water surface in laboratory vessels were shed there by ephippial females when moulting (despite the attendant risk of exposure to UV radiation). This implies that the moulting behaviour of female Daphnia may determine the fate of their dormant offspring, predetermining whether they remain in the natal environment (when the ephippium is released into the water column) or disperse (when it is deposited at the water surface).
• 4 Our findings reveal a potential mechanism underlying the high dispersal capacity of freshwater cladocerans inhabiting island‐like aquatic habitats.

     

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.     

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.     

Side effects of water liming on body size and eggs development of Daphnia species in Lake Orta (N. Italy)

Life-tables of the Daphnia species (D. longispina and D. obtusa), currently inhabiting the previously acidified Lake Orta, were followed to study the side effects of liming on the demographic parameters (e.g. longevity, body size increment, eggs and new-borns production), of both species. Different media were prepared to simulate the lake water pH's (5, 6 and 7), and copper concentrations (40 μg l^-1), before and after liming, using GF/C filtered lake water in such simulation and as a reference medium. The obtained results have shown that, the individuals of D. longispina were only able to survive after the liming and the consequent reduction in copper concentrations. However, due to shift occurred in the different media pH's after liming toward slightly alkaline conditions, no significant variations in body size increment or in eggs and newborns production were detected of this species in the different media (p = 0.38, p = 0.29, p = 0.09 respectively). On the other hand, the effect of the variations in pH on the body size increment of the B. obtusa individuals was marginally significant (p = 0.07). A highly significant (p = 0.002), increase in eggs production by D. obtusa individuals was detected in the different media after the CaCO₃ addition. Conversely, a corresponding highly significant (p = 0.0007), decrease in the production of new-borns, accompanied by the degeneration of about 78% of the produced eggs, was detected in the different media after liming. Our results suggest that in Lake Orta the actual irregular abundance of the studied Daphnia species and the eggs degeneration phenomenon, observed in our study, could be largely attributed to the interspecific competition resulting from species-specific sensitivity to the continuous variations in the lake water Chemistry after the liming operation. This revised version was published online in July 2006 with corrections to the Cover Date.     

Effects of winter conditions on Daphnia dynamics and genetic diversity in a dimictic temperate reservoir

1. Winter conditions shape plankton dynamics and community composition in temperate regions, but their effect on dynamics and genetic composition of cyclical parthenogens like Daphnia is largely unclear. 2. For 5 years, we studied the dynamics, hatching from resting eggs and genetic structure of a D. galeata × longispina hybrid complex in a dimictic, temperate reservoir. Our main hypothesis was that higher spring densities and an earlier population peak will be observed after warmer winters, with a lower genetic diversity because of a lower contribution of resting eggs to population growth. 3. The study period could clearly be categorised into cold‐winter years (n = 3) and warm‐winter years (n = 2). Daphnia densities at the end of spring overturn were ～10‐fold lower after cold winters than after warm ones, but no pattern emerged concerning the timing and the height of the population peak in early summer. 4. Hatching intensity from resting eggs was higher and contributed up to 8.5% to Daphnia abundance in a cold‐winter year compared to a negligible contribution in a warm‐winter year. Consistent with this finding, new multilocus genotypes (MLGs) adding to the overwintering stock after the end of spring overturn and presumably originating from resting eggs increased genetic diversity and attained high frequencies within the population only after a cold winter. New MLGs were recorded also after warm winters, but they never gained dominance and no shift in genetic diversity was observed. However, genetic diversity was not generally reduced after warm winters. 5. Our results confirm earlier findings that winter conditions have only a limited effect on the main growth phase and the peak of Daphnia during late spring and early summer. However, winter conditions determine the contribution of resting eggs to the population development, which may profoundly alter the genetic composition of the population compared to the previous season.     

Low crowding threshold for induction of sexual reproduction and diapause in a Patagonian rotifer

1. This study investigates the basis and ecological significance of the extremely high propensity for mixis (sexual reproduction) observed in laboratory populations of Brachionus calyciflorus from a temporary pond in Patagonia. 2. Experiments with stem females hatched from resting eggs showed that these females were exclusively amictic but produced mictic daughters even at very low population densities. In six experiments, newly hatched stem females were cultured individually in different volumes (1.5, 12, 40 and 150 mL). The percentage of mictic daughters (mixis ratio) was high in the smaller volumes (e.g. 44–83% in 1.5 mL) and lower in the largest volume (6–21% in 150 mL). A regression analysis combining the data from these experiments showed a highly significant decrease in mixis ratio with volume and indicated that the lowest population density at which mixis still occurs (mixis threshold) would be 3.4 females L^?1 (95% CL 2.9–4.0 females L^?1). This value is considerably lower than mixis thresholds for other rotifers (25–250 females L^?1 for many species and 9000–477 000 females L^?1 for some Spanish Brachionus plicatilis). 3. In three additional experiments, stem females and their amictic daughters were cultured individually in 150 mL. The percentage of mictic daughters produced by these two generations of females was not significantly different, showing that the mixis response to crowding is not inhibited in the stem‐female generation. 4. Laboratory experiments showed that two common predators of the Patagonian B. calyciflorus (the calanoid copepod Parabroteas sarsi and the backswimmer Notonecta vereertbruggheni) each ate 60–70 B. calyciflorus predator^?1 per day and cleared all rotifers from c. 250 mL per day. Thus, a very low mixis threshold and high maximal mixis ratio may ensure production of some resting eggs soon after colonisation of the pond and before complete removal from the plankton. 5. Two laboratory experiments showed that resting eggs of the Patagonian B. calyciflorus hatched at variable rates (28 and 81%) after a brief diapause when kept in the conditions under which they were produced and oviposited (20–21 °C; L: D 16: 8). Early hatching of resting eggs from pond sediment may allow multiple periods of colonisation and resting egg production in a season. This may offset the fitness cost of limited population growth through female parthenogenesis in the face of unpredictable and abrupt risk of extinction because of predators.     

Finding the optimal vertical distribution: behavioural responses of Daphnia pulicaria to gradients of environmental factors and the presence of fish

1. The vertical distribution of zooplankton results from active habitat choice aiming to optimise fitness gain in a system of trade‐offs. 2. Using large, controlled indoor mesocosms (Plön Plankton Towers), we monitored the behavioural response of Daphnia pulicaria to vertical gradients of temperature, food, oxygen and light, in the presence and absence of fish predation. 3. In the absence of fish, Daphnia distributed as predicted by an ideal ‘free distribution with costs’. If the food was distributed homogeneously, they stayed in the warm epilimnion, while they balanced their time dwelling in epi‐ and hypolimnion if the food was concentrated in a deep‐water maximum. 4. However, oxygen depletion in the hypolimnion, representing an additional cost, prevented Daphnia from completely exploiting the hypolimnetic food maximum. Consequently, the proportion dwelling in the hypolimnion was larger if oxygen was not limiting. 5. Fish predation had an overwhelming effect, driving Daphnia into the hypolimnion under all experimental conditions. If permitted by oxygen availability, Daphnia used the whole hypolimnion, but oxygen depletion reduced their possible habitat to the upper hypolimnion with oxygen concentrations above c. 0.7 mg L^?1. As fish were less tolerant of low oxygen, the layer below the thermocline formed a predation refuge for Daphnia.     

Breeding Daphnia magna

This paper deals with some methods for breeding Daphnia magna. Critical variables are the filtration of the medium on charcoal, the composition of the medium, the amount of food, the addition of vitamins with yeast extract and the extent of refreshing the medium. If these variables are well controlled, breeding daphnia is easy with reproduction varying from 3 to 9 young per daphnia per day.     

A major shift in Daphnia genetic structure after the first ice‐free winter in a German reservoir

1. Climate warming may cause disruption of trophic linkages in aquatic ecosystems and lead to changes in abundance and genetic structure of zooplankton populations. We monitored the community of the Daphnia galeata‐hyalina hybrid complex in the Saidenbach Reservoir (Saxony, Germany) using allozyme electrophoresis for three consecutive years (2005–07), including one (2007) following an unusually warm winter that prevented the formation of ice cover for the first time in the history of the reservoir. 2. Genetic composition during the 2007 season differed substantially from the two preceding years that experienced the usual 3‐month ice period. Three abundance peaks in June, July and October 2007 were dominated by hybrids of Daphnia galeata x hyalina, whereas in the 2005 and 2006 seasons two peaks in June and September were dominated by Daphnia hyalina genotypes. 3. The genetic composition of the pool of diapausing eggs produced in autumn and the rate of change of genotype abundance during the following spring indicate recruitment of the D. hyalina subpopulation from ex‐ephippial animals during the spring population increase. 4. The differing potential to contribute to the overwintering animal pool or to the inoculum from diapausing eggs was confirmed by results from laboratory life‐table experiments. Daphnia galeata clones survived longer and produced parthenogenetic offspring under winter conditions, whereas D. hyalina clones showed a shorter lifespan and produced resting eggs. 5. Our results indicate a profound role of recruitment strategy in the observed shift in genetic composition. Increasing winter temperatures predicted in the context of climate change may thus favour overwintering animals, leading to an increase in the contribution of these genotypes to the population. Such microevolutionary processes may dampen possible seasonal mismatches between daphnid populations and their food or predator populations.