Circadian Rhythm in Feeding Behavior of Daphnia dentifera

Circadian rhythms enable organisms to mediate their molecular and physiological processes with changes in their environment. Although feeding behavior directly affects within-organism processes, there are few examples of a circadian rhythm in this key behavior. Here, we show that Daphnia have a nocturnal circadian rhythm in feeding behavior that corresponds with their diel vertical migration (DVM), an important life history strategy for predator and UV avoidance. In addition, this feeding rhythm appears to be temperature compensated, which suggests that feeding behavior is robust to seasonal changes in water temperature. A circadian rhythm in feeding behavior can impact energetically demanding processes like metabolism and immunity, which may have drastic effects on susceptibility to disease, starvation risk, and ultimately, fitness.

Density, parasitism, and sexual reproduction are strongly correlated in lake Daphnia populations

Many organisms can reproduce both asexually and sexually. For cyclical parthenogens, periods of asexual reproduction are punctuated by bouts of sexual reproduction, and the shift from asexual to sexual reproduction has large impacts on fitness and population dynamics. We studied populations of Daphnia dentifera to determine the amount of investment in sexual reproduction as well as the factors associated with variation in investment in sex. To do so, we tracked host density, infections by nine different parasites, and sexual reproduction in 15 lake populations of D. dentifera for three years. Sexual reproduction was seasonal, with male and ephippial female production beginning as early as late September and generally increasing through November. However, there was substantial variation in the prevalence of sexual individuals across populations, with some populations remaining entirely asexual throughout the study period and others shifting almost entirely to sexual females and males. We found strong relationships between density, prevalence of infection, parasite species richness, and sexual reproduction in these populations. However, strong collinearity between density, parasitism, and sexual reproduction means that further work will be required to disentangle the causal mechanisms underlying these relationships.

Multiple factors likely explain variation in investment in sexual reproduction by lake Daphnia populations

Many organisms can reproduce both asexually and sexually. For cyclical parthenogens, periods of asexual reproduction are punctuated by bouts of sexual reproduction, and the timing of the shift from asexual to sexual reproduction has large impacts on fitness and population dynamics. We studied populations of Daphnia dentifera to determine the amount of investment in sexual reproduction as well as the factors associated with variation in investment in sex. To do so, we tracked host density, parasite infections, sexual reproduction, temperature, and light attenuation in 15 lake populations of D. dentifera for three years. We monitored infections by nine common parasites; this is notable since most prior studies on investment in sex and parasitism have focused on a single parasite, even though multiparasite communities are the norm in nature. We found substantial variation in investment in sex, with some populations reproducing entirely asexually throughout the study period and others shifting almost entirely to sexual reproduction by late autumn. We found that higher host density and parasitism were associated with greater investment in sex. Temperature and light attenuation were not as predictive of investment in sex, but received some statistical support. While correlational, our results leverage a large time series dataset and suggest multiple factors likely drive variation in sexual reproduction in this dominant member of lake food webs.

The effects of parasite exposure on mortality from aquatic contaminants, carbaryl and elevated salinity, in a freshwater crustacean

Freshwater pollution is a major global concern. Common methods for determining the effects of contaminants on freshwater organisms involve short-term laboratory experiments with otherwise healthy organisms. However, in natural systems, organisms are commonly exposed to parasites, which could alter their ability to survive exposure to aquatic contamination. We used a freshwater crustacean (Daphnia dentifera) to quantify the effects of parasite exposure on mortality from two common freshwater contaminants (elevated salinity [NaCl] and carbaryl). In our salinity trial, both parasite exposure and elevated salinity reduced survival in an additive manner. In our carbaryl trial, exposure to carbaryl reduced survival and we found a less-than-additive (i.e. antagonistic) interaction between carbaryl and the parasite; the parasite only reduced survival in the control (no carbaryl) treatments. Our results demonstrate that parasites and contaminants can jointly affect mortality in aquatic organisms in an additive or less-than-additive manner.

Genotypic variation in parasite avoidance behaviour and other mechanistic, nonlinear components of transmission

Traditional epidemiological models assume that transmission increases proportionally to the density of parasites. However, empirical data frequently contradict this assumption. General yet mechanistic models can explain why transmission depends nonlinearly on parasite density and thereby identify potential defensive strategies of hosts. For example, hosts could decrease their exposure rates at higher parasite densities (via behavioural avoidance) or decrease their per-parasite susceptibility when encountering more parasites (e.g. via stronger immune responses). To illustrate, we fitted mechanistic transmission models to 19 genotypes of Daphnia dentifera hosts over gradients of the trophically acquired parasite, Metschnikowia bicuspidata . Exposure rate (foraging, F ) frequently decreased with parasite density ( Z ), and per-parasite susceptibility ( U ) frequently decreased with parasite encounters ( F × Z ). Consequently, infection rates ( F × U × Z ) often peaked at intermediate parasite densities. Moreover, host genotypes varied substantially in these responses. Exposure rates remained constant for some genotypes but decreased sensitively with parasite density for others (up to 78%). Furthermore, genotypes with more sensitive foraging/exposure also foraged faster in the absence of parasites (suggesting ‘fast and sensitive’ versus ‘slow and steady’ strategies). These relationships suggest that high densities of parasites can inhibit transmission by decreasing exposure rates and/or per-parasite susceptibility, and identify several intriguing axes for the evolution of host defence.

Toxins or medicines? Phytoplankton diets mediate host and parasite fitness in a freshwater system

Diets must satisfy the everyday metabolic requirements of organisms and can also serve as medicines to combat disease. Currently, the medicinal role of diets is much better understood in terrestrial than in aquatic ecosystems. This is surprising because phytoplankton species synthesize secondary metabolites with known antimicrobial properties. Here, we investigated the medicinal properties of phytoplankton (including toxin-producing cyanobacteria) against parasites of the dominant freshwater herbivore, Daphnia. We fed Daphnia dentifera on green algae and toxic cyanobacteria diets known to vary in their nutritional quality and toxin production, and an additional diet of Microcystis with added pure microcystin-LR. We then exposed Daphnia to fungal and bacterial parasites. Anabaena , Microcystis and Chlorella diets prevented infection of Daphnia by the fungal parasite Metschnikowia , while Nodularia toxins increased offspring production by infected hosts. In contrast to their medicinal effects against Metschnikowia , toxic phytoplankton generally decreased the fitness of Daphnia infected with the bacterial parasite, Pasteuria . We also measured the amount of toxin produced by phytoplankton over time. Concentrations of anatoxin-a produced by Anabaena increased in the presence of Metschnikowia , suggesting parasite-induced toxin production. Our research illustrates that phytoplankton can serve as toxins or medicines for their consumers, depending upon the identity of their parasites.