Many representatives of freshwater zooplankton produce at some stage in their life cycle resting stages. A variable portion of the eggs of the previous growing period will hatch at the next occasion while the remaining ones are added to a persistent egg bank, where they can remain viable for decades or longer. The importance of the study of resting eggs and egg banks in general for such different disciplines as taxonomy, ecological biogeography, paleolimnology, nature conservation, evolutionary ecology and community and population ecology is generally appreciated. The major current and expected future developments in this rapidly expanding field of research are presented here. The structure and dynamics of the egg bank are determined by the life history characteristics of the species (or local population), the hatching phenology of their resting stages, and the characteristics of the habitat. The horizontal distribution of dormant stages is generally patchy, with a greater density in the deeper and/or windward parts of a pond or lake. In sediment cores, most viable (responsive) eggs occur in the upper centimeters, although vertical variation related to the history of fish predation or water quality occurs. The accumulation of resting stages of different species, generations and genotypes with variable regeneration niches results in a mixed egg bank with greater potential biodiversity than the active community sampled at any one moment. Through the benthic–pelagic coupling, this dormant reservoir may have considerable impact on the evolutionary potential of the organisms, the ecological dynamics of the community and the distribution of species. Egg banks can be considered the archive of the local habitat, since the pattern of changes in species assemblage and genotypes from the past up to the present reflect changes due to natural or anthropogenic impact that can be used to reconstruct evolutionary processes or even to restore the local habitat. Overlooking the egg bank as an important component of zooplankton communities may lead to erroneous interpretations in the analysis of community and population genetic structure. This review integrates technical and scientific information needed in the study of the structure and function of egg banks in zooplankton with special focus on the fascinating latest developments in the field.

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Egg banks in freshwater zooplankton: evolutionary and ecological archives in the sediment

The factors inducing sexual eggs, as well as the offspring sex ratio preceding sexual egg formation, were investigated in a cyclic parthenogen, Daphnia magna (Crustacea, Cladocera). Laboratory experiments were conducted on individual animals living in low-through chambers, making it possible to separate the effects of two density-dependent factors: food limitation and a chemically mediated cue. These factors were studied under a short-day photoperiod and in permanent light. The simultaneous actions of an inductive photoperiod, food limitation and chemically mediated crowding, were all needed to induce sexual egg formation. When only two stimuli were present, the offspring sex ratio was 0.50 or lower, and no sexual eggs were produced

Sexual reproduction in Daphnia magna requires three stimuli

For a short-lived organism, such as a freshwater zooplankter, the ways of coping with years of local recruitment failure are either to disperse between habitats and recolonise or to disperse in time through diapause. Diapause is common among freshwater zooplankton and is generally seen as a way to escape periods of harsh environmental conditions. The egg-bank or pool of diapausing copepodites in lake sediments resulting from the production of diapausing stages has several implications for zooplankton ecology, genetics, and evolution which we outline in this review. The presence of a benthic dormant stage also creates a coupling between the benthic habitat and the pelagic, and we argue that zooplankton phenology is a result of selective forces in both habitats. The spatial distribution of diapausing eggs appears to be governed by random resuspension dynamics coupled with higher hatching rates in shallow waters. For diapausing copepodites, however, an active choice of where and how deep to enter the sediment may affect their distribution. In a reanalysis of published data, we found a size-dependent bathymetric distribution and vertical distribution in the sediment of diapausing cyclopoid copepodites. Our review of published laboratory studies showed that predictors of seasonal change such as photoperiod and temperature were the only type of cues used for the termination of diapause. We also found a relation between generation length and the type of cue used for diapause induction: copepods mainly used seasonal cues from the abiota, rotifers mainly used cues from the biotic environment, and cladocerans used a mix of both types. We describe patterns in emergence timing and contribution to population dynamics from studies using in situ estimation of emergence, and conclude that hatching from dormant stages may qualitatively and quantitatively affect zooplankton population dynamics and seasonal succession.

Dormancy in freshwater zooplankton: Induction, termination and the importance of benthic-pelagic coupling

The factors inducing male offspring and the mechanism of sex-ratio adjust- ment were investigated in Daphnia magna, a cyclically parthenogenetic cladoceran (Crus- tacea). Laboratory experiments were conducted on individual animals living in flow-through chambers, to separate the effects of various density-dependent factors. Fluctuating food levels had no effects on offspring sex when starvation was not severe. Water from crowded populations of different Daphnia species contained substances that induced up to 43% male broods. The chemically mediated crowding effect was detectable under conditions repre- senting a low population density, implying that animals responded to their own metabolites. Daphnia species differed in their inducing effects, and in one case (water from D. pulex) strong male offspring induction was associated with growth inhibition. A short daylength was also demonstrated to induce male offspring effectively. Evidence was found to support a mechanism of sex-ratio regulation that implies a regular alternation of brood sex, resulting in a sex ratio close to 1/2, the expected optimum ratio. Even individual mothers adjusted the sex ratio of their offspring within a sequence of seven broods. No resting eggs were formed in the experiments, indicating that formation of male offspring and resting eggs are independently controlled, possibly by distinct sets of environmental cues.

Sex Determination in Daphnia Magna

SUMMARY. A method is described whereby ephippia (sexual eggs plus protective membranes) are experimentally induced at low food levels (≤ 0.05 mg spinach D. magna-1), high female culture-densities (≤ 0.4 D. magna ml-1) and in short-day photoperiods (L:D < 12:12, 12°C). The density-dependent ephippial response was related to an increased encounter rate between females at the higher densities. External metabolites had no significant effect on ephippia production. Ephippia were formed in the second generation after exposure to short day-lengths. Five clones from the same population exhibited genetic variation in their critical photoperiods for induction of sexual reproduction (≥= 10% ephippia). There was evidence of a genetic difference in photoresponse between two populations of D. magna. The evolution of timing mechanisms for ephippia production are discussed in relation to the predictability of environmental change.

The effect of food availability, female culture-density and photoperiod on ephippia production in Daphnia magna Straus (Crustacea: Cladocera)

1. Control of the sexual polymorphism leading to an embryonic diapause was studied in an arctic (71° N) population of Daphnia middendorffiana. The reproductive cycle is similar to a previously analyzed population of D. pulex although males are non-functional in the polymorphism.2. In constant temperatures and standardized culture density the reproductive shift is controlled by daylength with a critical photoperiod of L:D 22:2 at 12° C. Control may be partially overridden in long days (constant light) by density of culture. The overwintering generation shows resistance to both photoperiodic induction and density suppression, and one or two non-diapause broods are released before reproduction shifts.3. The reproductive shift in the source pool at Barrow, Alaska occurs in mid-July when the sun is continuously above the horizon and after the overwintering generation has released one brood of non-diapausing embryos. Completeness of the shift and the relatively low density of the population in the pool argues a...

Raymond G. Stross

Papers

Photoperiod control of diapause in daphnia. ii. induction of winter diapause in the arctic