Is Survival After Pathogen Exposure Explained by Host's Immune Strength? A Test with Two Species of White Grubs (Coleoptera: Scarabaeidae) Exposed to Fungal Infection

A major innate defense system in invertebrates is the melanization of pathogens and damaged tissues. This important process is controlled by the enzyme phenoloxidase (PO) that in turn is regulated in a highly elaborate manner for avoiding unnecessary production of highly toxic and reactive compounds. Recent progress, especially in arthropods, in the elucidation of mechanisms controlling the activation of zymogenic proPO into active PO by a cascade of serine proteinases and other factors is reviewed. The proPO-activating system (proPO system) is triggered by the presence of minute amounts of compounds of microbial origins, such as beta-1,3-glucans, lipopolysaccharides, and peptidoglycans, which ensures that the system will become active in the presence of potential pathogens. The presence of specific proteinase inhibitors prevents superfluous activation. Concomitant with proPO activation, many other immune reactions will be produced, such as the generation of factors with anti-microbial, cytotoxic, opsonic, or encapsulation-promoting activities.

Evolutionary ecology seeks to understand the selective reasons for the design features of the immune defense, especially with respect to parasitism. The molecular processes thereby set limitations, such as the failure to recognize an antigen, response specificity, the cost of defense, and the risk of autoimmunity. Sex, resource availability, and interference by parasites also affect a response. In turn, the defense repertoire consists of different kinds of immune responses--constitutive or induced, general or specific--and involves memory and lasting protection. Because the situation often defies intuition, mathematical analysis is typically required to identify the costs and benefits of variation in design, but such studies are few. In all, insect immune defense is much more similar to that of vertebrates than previously thought. In addition, the field is now rapidly becoming revolutionized by molecular data and methods that allow unprecedented access to study evolution in action.

The evolutionary-ecology approach to studying immune defences has generated a number of hypotheses that help to explain the observed variance in responses. Here, selected topics are reviewed in an attempt to identify the common problems, connections and generalities of the approach. In particular, the cost of immune defence, response specificity, sexual selection, neighbourhood effects and questions of optimal defence portfolios are discussed. While these questions still warrant further investigation, future challenges are the development of synthetic concepts for vertebrate and invertebrate systems and also of the theory that predicts immune responses based on a priori principles of evolutionary ecology.