Non-immunological defense in an evolutionary framework.

After parasite infection, invertebrates activate immune system-based defenses such as encapsulation and the signaling pathways of the innate immune system. However, hosts are often able to defend against parasites without using these mechanisms. The non-immunological defenses, such as behaviors that prevent or combat infection, symbiont-mediated defense, and fecundity compensation, are often ignored but can be important in host-parasite dynamics. We review recent studies showing that heritable variation in these traits exists among individuals, and that they are costly to activate and maintain. We also discuss findings from genome annotation and expression studies to show how immune system-based and non-immunological defenses interact. Placing these studies into an evolutionary framework emphasizes their importance for future studies of host-parasite coevolution.