Effects of juvenile host density and food availability on adult immune response, parasite resistance and virulence in a Daphnia-parasite system

Host density can increase infection rates and reduce host fitness as increasing population density enhances the risk of becoming infected either through increased encounter rate or because host condition may decline. Conceivably, potential hosts could take high host density as a cue to up-regulate their defence systems. However, as host density usually covaries with food availability, it is difficult to examine the importance of host density in isolation. Thus, we performed two full-factorial experiments that varied juvenile densities of Daphnia magna (a freshwater crustacean) and food availability independently. We also included a simulated high-density treatment, where juvenile experimental animals were kept in filtered media that previously maintained Daphnia at high-density. Upon reaching adulthood, we exposed the Daphnia to their sterilizing bacterial parasite, Pasteuria ramosa, and examined how the juvenile treatments influenced the likelihood and severity of infection (Experiment I) and host immune investment (Experiment II). Neither juvenile density nor food treatments affected the likelihood of infection; however, well-fed hosts that were well-fed as juveniles produced more offspring prior to sterilization than their less well-fed counterparts. By contrast, parasite growth was independent of host juvenile resources or host density. Parasite-exposed hosts had a greater number of circulating haemocytes than controls (i.e., there was a cellular immune response), but the magnitude of immune response was not mediated by food availability or host density. These results suggest that density dependent effects on disease arise primarily through correlated changes in food availability: low food could limit parasitism and potentially curtail epidemics by reducing both the host's and parasite's reproduction as both depend on the same food

Testing small clutch size models with Daphnia.

Life-history theory predicts that for small clutches, variance in egg size (between individuals) should decrease in a predictable invariant manner as clutch size increases. To test this, we studied Daphnia magna at 350 different food treatments and recorded the number of eggs and the volume of each egg for their first clutch. As predicted, we found that the relationship between clutch size and resources devoted to reproduction was linear, variance in egg volume decreased with increasing clutch size, and resources were shared relatively equally between the eggs in a clutch. However, we found that the rate at which the range of egg volumes decreased with clutch size was slower than predicted. We discuss possible explanations for this discrepancy, including a lower limit on the volume of eggs that are produced and selection for smaller eggs when food is abundant. Consistent with this, we found that mean egg volume decreased with increasing clutch size

Cooperation and the scale of competition in humans.

Explaining cooperation is one of the greatest challenges for evolutionary biology. It is particularly a problem in species such as humans, where there is cooperation between nonrelatives. Numerous possible solutions have been suggested for the problem of cooperation between nonrelatives, including punishment, policing, and various forms of reciprocity. Here, we suggest that local competition for resources can pose a problem for these hypotheses, analogous to how it can select against cooperation between relatives. We extend the prisoner's dilemma (PD) game to show that local competition between interacting individuals can reduce selection for cooperation between nonrelatives. This is because, with local competition, fitness is relative to social partners, and cooperation benefits social partners. We then test whether nonrelated humans adjust their level of cooperation facultatively in response to the scale of competition when playing the PD for cash prizes. As predicted, we found that individuals were less likely to cooperate when competition was relatively local. Cooperation between humans will therefore be most likely when repeated interactions take place on a local scale between small numbers of people, and competition for resources takes place on a more global scale among large numbers of people