Daphnia exhibit diurnal vertical migration in shallow rock-pools

We report diurnal changes in vertical distribution ofDaphnia longispina andD. magna in shallow (25–60 cm) rock-pools on Baltic islands, southern Finland. During the daytime shallow rock-poolDaphnia reside in water layers close to the sediment while at midnight they either are evenly distributed throughout the water column, or have a modal peak of distribution in the surface water. Contrary to this pattern, no changes were observed in vertical distribution ofD. longispina at noon and at midnight in a 150 cm deep pond. We suggest that the pattern ofDaphina vertical migration in shallow rock-pools is not satisfactorily explained by predation.     

Content and turnover rates of phosphorus in Daphnia pulex: Effect of food quality

The content of phosphorus in individual Daphnia pulex and the rates at which the element turns over are functions of the P contents of algal cells fed to the zooplankton. Chlamydomonas and Ankistrodesmus were grown in semi-continuous cultures containing 2 µM and 10 µM PO₄ to produce differences in cellular P of the algae. Animals reared on the cultures high in P contained 60 percent more P than did animals of equal size that were fed low P cells. Daphnia with high body contents of P turned over that P faster than did the animals fed low P algae. Measured turnover rates imply that large differences exists between rates of gross P assimilation and P excretion for animals fed high and low P sources.     

Diel patterns of feeding and vertical migration in daphnids and diaptomids during the clear water phase in Lake Geneva (France)

In lacustrine environments, little attention has been paid to small-scale interactions between zooplankton diel vertical migration (DVM) and feeding rhythms. Moreover, most of the information on in situ diel feeding and migratory rhythms is based on low sampling frequencies. The kinetics and the degree of coupling of these processes are thus only roughly known. Here, we present a study conducted on a diel cycle in Lake Geneva to establish the temporal and spatial relationships between DVM and grazing activity of the dominant planktonic crustaceans. Our methodological approach is based on reliable and frequent (every 30 minutes) sampling, and on gut fullness analysis. We test the hypothesis of temporal and spatial segregation in DVM and feeding activity of sympatric taxa to counteract resource competition. We also evaluate the variation in DVM and feeding activity between taxa, size and sexes. In Lake Geneva, the Daphnia complex of different species and size (D. hyalina × galeata) and the diaptomid (Eudiaptomus gracilis) have distinct DVM and diel feeding patterns which lead to temporal and spatial segregation. Differences arise from the amplitude and kinetics of DVM and diel feeding rhythms. A strong day/night contrast in depth distribution and feeding activity was observed for the large daphnids while the small daphnids and the diaptomids had lower amplitudes of DVM and weaker diel changes in feeding activity. Large Daphnia exhibited a bimodal feeding pattern coupled with dynamic interchange of individuals between the epi- and hypolimnetic layers at dusk and dawn. In contrast, little coupling between DVM and feeding patterns was found for the diaptomid. These distinct behaviours can be viewed as specific adaptive strategies developed by calanids and daphnids to limit interspecific competition and to compromise between avoidance of starvation in deep waters and avoidance of visual predators in surface layers. Our study supports the hypothesis of exogenous control of Daphnia DVM by the relative change in light intensity at dusk and dawn, but also suggests that small Daphnia (not large ones), are controlled by absolute light variations when this major stimulus is lacking. Our results also support the hypothesis that selective predation by fish is responsible for the observed differences in DVM and diel feeding patterns of sized-daphnids and diaptomids. Other factors explaining the coupling of DVM and feeding patterns are hunger, vertical temperature gradient and for daphnids, size. Thus, ecological plasticity in crustacean DVM and feeding patterns results from the interactive effect of multiple abiotic and biotic driving forces. Finally, our study also shows that large Daphnia have a marked contribution to the acceleration of downward nutrient fluxes in Lake Geneva, via their diurnal rhythm in feeding and vertical migration. Ecological implications of the study for lake management and sampling design of zooplankton grazing studies are also presented.

---

Résumé Dans les écosystèmes lacustres, les interactions à fine échelle temporelle entre les patrons diurnes de migration verticale et de broutage du zooplancton sont peu étudiées. En outre, jusq'à présent, les études ont généralement été réalisées selon des chroniques temporelles assez lâches. La cinétique et le degré d'interaction entre les patrons journaliers de migration verticale et de broutage sont donc encore mal connus. La présente étude, conduite au Lac Léman (Lac de Genéve) au cours d'un cycle nycthéméral, tente de préciser les liaisons spatiales et temporelles existant entre les migrations journalières et la consommation de phytoplancton chez les taxons de Crustacés les mieux représentés. Notre approche méthodologique repose sur une maille temporelle d'échantillonnage fine et sur l'analyse de la fluorescence du contenu stomacal. Nous testons l'hypothèse d'une ségrégation spatio-temporelle visant à réduire la compétition entre les taxons sympatriques et reposant sur des différences entre les patrons journaliers respectifs de migration et d'alimentation. Nous évaluons pour ces rhythmes d'activité les différences entre les espèces, les classes de tailles et les sexes. Au lac Léman, le complexe de différentes espèces et tailles de daphnies (Daphnia hyalina × galeata) et le diaptomide (Eudiaptomus gracilis) présentent des patrons journaliers de migration et d'alimentation distincts, assurant une ségrégation spatio-temporelle. Les différences proviennent de variations dans la cinétique et l'amplitude des migrations et dans les niveaux d'alimentation. Les grandes daphnies affichent un fort contraste jour/nuit dans leur répartition verticale et leur état de réplétion, tandis que les petites daphnies et les diaptomides présentent une faible amplitude de migration et de variations circadiennes de réplétion. Les grandes daphnies ont un rythme alimentaire bimodal couplé avec un relais dynamique des organismes entre l'épilimnion et l'hypolimnion au crépuscule et à l'aube. Chez le diaptomide, les interactions sont au contraire faibles entre les patrons de migration verticale et de réplétion. Ces différents comportements peuvent être perçus comme des stratégies adaptives spécifiques développées par les daphnies et les diaptomides pour limiter la compétition interspécifique et aboutir à un compromis satisfaisant entre l'évitement de la famine en eaux profondes et de la prédation par les poissons dans les eaux superficielles. Notre étude conforte l'hypothèse d'un contrôle exogène de la migration verticale de Daphnia par les changements relatifs de la lumière au crépuscule et à l'aube. En l'absence de ce stimulus, la répartition verticale des petites daphnies semble par contre contrôlée par les variations absolues de lumière. Nos observations confortent également l'hypothèse que la prédation sélective par les poissons est responsable des différences observées dans les patrons de migration des grandes daphnies et ceux des petites daphnies et des diaptomides. Les autres facteurs pouvant influencer les patrons de migration et d'alimentation des crustacés du Lac Léman sont la famine, le gradient thermique vertical et, chez les daphnies, la taille. En définitive, la plasticité écologique des patrons journaliers de migration et d'alimentation résulte des effets interactifs de daphnies ont un rôle très important dans le transfert des nutriments dans les couches profondes durant l'été, via leurs migrations verticales et les variations circadiennes d'activité alimentaire. Les implications écologiques pour l'aménagement lacustre et la planification des études portant sur le broutage du zooplancton sont aussi présentées.

     

Contrasting migration behaviour of Daphnia pulicaria and D. galeata x hyalina, in avoidance of predation by 0+perch (Perca fluviatilis)

In spring and early summer, a small population of the large-bodied Daphnia pulicaria coexists with a much larger population of the medium-sized hybrid Daphnia galeata × hyalina in the epilimnion of Lake Maarsseveen (The Netherlands). When large shoals of juvenile perch (Perca fluviatilis) appear in the open water, both species start to migrate vertically. Since D. pulicaria has a larger body-size than D. galeata × hyalina, and is therefore competitive dominant over the hybrid, it is unlikely that both species interact via their common food resource, but they react both to fish predation. However, since they differ in size, and therefore in vulnerability for fish predation, both species adopt different strategies. The smaller bodied, and less vulnerable D. galeata × hyalina exhibits diel vertical migration ascending to the surface at dusk, and staying there during the night. In this way, benefiting from the higher temperatures of the surface layers. In contrast, the large-bodied, and more vulnerable D. pulicaria selects the deep cold hypolimnion water layers as refuge against fish predation. In this way it benefits from a safe habitat, free from fish predators, but on the other hand suffers from low water temperatures, which decrease its fitness. It is likely that the relatively higher temperature in the upper water layers serves as a proximate factor for the downward migration of D. pulicaria.     

Daphnia vertical distribution and the presence of toxic cyanobacteria

Daphnia can alter its vertical position in the water column in response to chemical cues from predators. In this study we tested the hypothesis that a Daphnia pulex clone with little evolutionary exposure to cyanobacteria would move away from patches of cyanobacteria (Anabaena affinis and A. flos-aquae) which contain potent endotoxins. Daphnia was censused at 2 h intervals for 6 h in laboratory columns in which there was a steep vertical gradient of cyanobacteria. Data were analyzed by repeated-measures ANOVA. Control (no Anabaena) and experimental columns showed no significant differences in Daphnia distributions. Daphnia in experimental columns frequently moved into areas with dense concentrations of Anabaena and stayed there for long periods of time. Our results show that this D. pulex clone does not exhibit a rapid (within 6 h) avoidance response to toxic Anabaena.     

Caridina nilotica in Lake Victoria: abundance, biomass, and diel vertical migration

Caridina nilotica (Decapoda: Atyidae) in offshore waters of Lake Victoria were investigated with both day and night sampling over a period of two years. Offshore populations are mainly planktonic rather than benthic, and the animals exhibit diel vertical migrations into near-surface waters at night. These changes in diel abundance as well as the size-frequency distribution of the migrating shrimp suggest that the migratory behavior is in response to visual planktivory, because only the very smallest individuals (2–4 mm) remain in surface waters during the day. During October 1992, abundances were estimated both by vertical net sampling and by underwater video transect methods. Concordance was established between abundances estimated by the two methods. Only about 9% (night) to 14% (day) of the Caridina population appeared to be epibenthic. We suggest that the behavior of the animal is consistent with the hypothesis that it is not a strict detritivore as previously reported; rather it may engage in facultative planktivory, especially at night.     

Size-structured risk assessments govern Daphnia migration

One of the more fascinating phenomena in nature is animal mass migrations and in oceans and freshwaters, diel variations in depth distribution of zooplankton are a phenomenon that has intrigued scientists for more than a century. In our study, we show that zooplankton are able to assess the threat level of ultraviolet radiation and adjust their depth distribution to this level at a very fine tuned scale. Moreover, predation risk induces a size-structured depth separation, such that small individuals, which we show are less vulnerable to predation than larger, make a risk assessment and continue feeding in surface waters during day, offering a competitive release from down-migrating larger animals. Hence, we mechanistically show that such simple organisms as invertebrate zooplankton are able to make individual, size-specific decisions regarding how to compromise between threats from both predators and UV radiation, and adjust their diel migratory patterns accordingly.     

Ultrastructure of the epidermis of Meara stichopi (Platyhelminthes,Nemertodermatida) and associated extra-epidermal bacteria

This preliminary mechanistic model of normal swimming and phototactic behaviour in individual Daphnia was constructed using data and assumptions based on experiments and observations. Swimming under constant light intensity is characterized by short periods of upward movements alternating with equal periods of downward movements. Two oscillators are proposed that generate these phases in swimming. Unexpected shifts in depth, as observed in D. magna and D. longispina, are also present in the swimming of the computer daphnid and thus seem to be inherent to the underlying mechanism. As in real daphnids, during relative decreases in light intensity of low velocity, positive phototactic upward swimming is stepwise. With increasing velocity in the change in light, these steps disappear. When the model is triggered by a natural increase in light at dawn, a small downward movement results. Migration distance can be increased to commonly found depths of migrating Daphnia by the introduction of a fish exudate factor into the model, which enhances the phototactic response. Since attenuation of light in the water affects the phototactic swimming response, it also influences migration distance. The results of model calculations agree quite well with an empirical relationship between Secchi disc depth and amplitude of diel vertical migration in a number of lakes.     

Diel vertical migration and feeding rhythm of Daphnia longispina and Bosmina corexoni in Lake Toya, Hokkaido, Japan

Diel vertical migration (DVM) and diel feeding rhythm of two cladocerans, Daphnia longispina and Bosmina coregoni were investigated at the pelagic area of Lake Toya (Hokkaido, Japan) in May, August and October 1992. Both species performed nocturnal DVM. The amplitude of DVM, however, became smaller from May to October. Such seasonal variations in DVM could not be explained by light penetration and/or water temperature. The two species had a clear feeding rhythm; they fed at night in May and October but also after sunrise in August. These feeding rhythms appeared to be related to the light-dark cycle, but were not necessarily associated with their DVM. We suggest that the diel feeding rhythm and DVM are regulated independently by light cues.     

Density and size distribution of Daphnia populations at different fish predation levels

Density and size structure changes of natural daphniids populations were studied in enclosures with a different level of fish predation. Daphnia pulicaria was totaly removed in high predation variants, and the differences of the mean body length both in adults and juveniles are apparent between low predation enclosure and enclosure without fish. Daphnia galeata was replaced by D. magna and D. pulicaria in the enclosure without fish. The decline of densities and the substantial (30–50%) and fast (during 12 days) shift of mean body length both in adults and juveniles of D. galeata was induced by the fish (carp fry) introduction to the high predation enclosures.     

The lack of vertical migration in Daphnia: the effect of homogenously distributed food

Neither Daphnia hyalina, Daphnia cristata, nor Daphnia cucullata vertically migrated in Lake Mikoajskie and Lake Majcz. We suggest that even under strong fish predation pressure there is no reason to migrate when seston is relatively homogenously distributed throughout the water column.     

Chaoborus and fish-mediated influences on Daphnia longispina population structure,dynamics and life history strategies

Summary This study examined the long term effects of predation by larvae of the midge Chaoborus and simulated fish predation on experimental Daphnia longispina populations. Chaoborus predation, relative to fish predation, led to populations composed of larger individuals as a whole, larger egg-bearing individuals, and a larger primiparous instar. Daphnia retained helmets beyond the first instar in response to the presence of Chaoborus. Both types of predation, relative to predator-free controls, reduced prey population size and rates of increase, but increased population death rates. The reduction in population size due to predation led to increased resource availability for individuals remaining in the populations and increased individual fecundity in the predation treatments. The differences noted between the Chaoborus, fish, and control treatments increased with predation intensity.     

Diel vertical migration by males,females, copepodids and nauplii in a limnetic population of diaptomus (Copepoda)

Diel vertical migration (DVM) by the sexes and life history stages of the copepod Diaptomus novamexicanus (Herrick) was investigated during two summers in a subalpine lake (Castle Lake, California, USA). Migratory behavior was quantified repeatedly by 1300 and 21–2200 hr sampling in the epilimnion. Population density and vertical distribution were estimated routinely at 1300 hr and at 2200 hr on selected dates. Males had greater night to day biomass differentials and ratios than females. The sexes showed comparable DVM participation (% of total biomass migrating) and amplitudes of migration (distance of ascent). Copepodids displayed higher participation, night/day ratios and amplitudes than younger copepods, but adult participation differed only from that of nauplii. Active participation in high amplitude ascents thus increased with age and was greatest in adult males. Analysis suggests that such relative behaviors probably occur in most limnetic Diaptomus, and that the magnitude of ascent into the epilimnion is associated with daytime vertical distribution.     

Contrasting patterns of body size for Daphnia species that segregate by habitat

Summary We investigate how body size of two coexisting Daphnia species varies among 7 lakes that represent a gradient of predation risk. The two species segregate vertically in stratified lakes; D. galeata mendotae is typically smaller and more eplimnetic than D. pulicaria. The extent of vertical habitat partitioning, however, varies seasonally within and among lakes in apparent response to predation intensity by epilimnetic planktivorous fishes. Daphnia pulicaria uses the epilimnion at low levels of fish predation but is restricted to the hypolimnion under high fish predation, whereas D. galaeta mendotae always utilizes the epilimnion. The species display contrasting patterns of genetic variation in neonate size and size at maturity. D. pulicaria is larger in lakes with higher fish and Chaoborus densities whereas D. galeata mendotae is smaller. This contrast in body size in lakes with high predation is associated with greater habitat segregation in those lakes. In lakes with low predation risk, the two species are similar in body size at birth and maturity.Authorship order alphabetical     

Diel vertical migration ofEudiaptomus gracilis during a short summer period

Several aspects of a diel vertical migration (DVM) of adultEudiaptomus gracilis in Lake Maarsseveen (The Netherlands) are described. The period of DVM lasted from the end of May until the middle of August. On May 21, 1989, the population was found divided into a deep dwelling part and a part in the upper five meter. Large shoals of juvenile perch were observed in the open water for the first time. On June 7, the whole population was down below 10 m and concentrated in a zone of high chlorphyll-a concentrations. One week later, a regular DVM was performed. The amplitude of this migration gradually decreased towards the end of the migration period. The ascent in the evening and the descent in the morning took place after sunset and before sunrise, respectively. The movements coincided with high relative changes in light intensity. Population size increased rapidly during the period of DVM but decreased again before the end of this period.     

Diel vertical migration of zooplankton: optimum migrating schedule based on energy accumulation

Zooplankton perform diel vertical migration (DVM) to avoid predators at the upper water layer, but often stay in the upper water layer throughout the day seeking food in spite of the presence of predators. This difference in migrating behavior has been explained by differences in environmental conditions or genetic differences. We examined theoretically how nutritious conditions of zooplankton individuals relate to determining different migrating behavior. A simple optimization model, maximizing the population growth rate, demonstrates that zooplankton individuals change their migrating behavior depending on the amount of accumulated energy. Such energy accumulation and its investment in reproduction are repeated every reproductive cycle. Therefore, unless the reproductive cycle is synchronized among individuals, different migrating behaviors will be observed within a population even if no genetic differences exist. Our model demonstrates that such coexistence of the two migrating behaviors is possible in natural Daphnia populations, and suggests that internal conditions of zooplankton individuals may be important as a factor for determining migrating behavior of zooplankton. This revised version was published online in July 2006 with corrections to the Cover Date.     

The genetics of phototaxis in Daphnia magna: Existence of three phenotypes for vertical migration among parthenogenetic females

Using a cloning technique, we revealed the existence of three phototactically distinct types in Daphnia magna, viz. negatives, positives, and gipsies. The latter migrate continuously between a low and a high light intensity. The expression of this behaviour is genetically determined, although positives and negatives can reversibly switch into gipsies.It is proposed that two opposite genotypes exist, with the gipsy behaviour epigenetically induced. All juveniles behave more or less uniformly, irrespective of their origin. The phototactic responses develop rather late during ontogenesis.     

A hypothesis on the role of Aphanizomenon in translocating phosphorus

A mechanism for the upward translocation of phosphorus by flake-forming Aphanizomenon is hypothesized. Aphanizomenon germinates on oxic sediments and rises into the water column, then the flakes diurnally migrate. Observations from Spring Lake, Minnesota, U.S.A., indicate that Aphanizomenon translocates phosphorus to the summertime epilimnion. However, the observations cannot discriminate between which of two mechanisms related to Aphanizomenon flakes, upon germination or with subsequent diurnal migrations, is more important.     

The effect of different zooplankton grazing patterns resulting from diel vertical migration on phytoplankton growth and composition: a laboratory experiment

Diel vertical migration (DVM) of herbivorous zooplankton is a widespread behavioural phenomenon in freshwater ecosystems. So far only little attention has been paid to the impact of DVM on the phytoplankton community in the epilimnion. Some theoretical models predict that algal population growth in the epilimnion should depend on the herbivores migration and grazing patterns: even if migrating zooplankton consume the same total amount of algae per day in the epilimnion as non-migrating zooplankton, nocturnal grazing should result in enhanced algal growth and favour algal species with high intrinsic growth rates over species with lower intrinsic growth rates. To test these hypotheses we performed experiments in which several algal species were confronted with different feeding regimes of Daphnia. In the experiments algal growth did not only depend on the absolute time of grazing but was comparatively higher when grazing took place only during the night, even when the grazing pressure was the same. Furthermore, algal species with higher intrinsic growth rates had higher advantages when being grazed upon only discontinuously during the night than algal species with a smaller intrinsic growth rate. The grazing pattern itself was an important factor for relative algal performance.     

Intersex and male development in Daphnia magna

Sex induction is environmentally stimulated in Daphnia and involves a cue-dependent response for sex determination. Somatic growth was shown to be similar in males and females during juvenile instars, but divergent due to a reduction in male somatic growth, at about the time that females produce ovaries. At this time, males appeared morphologically adult with respect to secondary sex characteristics. Intersex was rare and is unlikely to be important in natural populations. Intersex could be induced in both sexes, and observed more frequently after longer exposure to high temperature, or in the second generation following a temperature change. This indicates an impact on the ability of mothers to determine the sex of the offspring. It may be possible to use intersex characteristics for manipulative investigation of sex-determination mechanisms in Daphnia. Although sex-determination is initiated before birth, intersex occurrence suggests that development of male characters requires an additional process, probably involving hormone activity, during juvenile development and maturation.