Six costs of immunity to gastrointestinal nematode infections.

The strength of the immune response and the outcome of the interaction of a host with a parasite are influenced by genetic and phenotypic characteristics of both parties, and by environmental variables. Allocation of host resources to immune defence reduces resources available for other life-history traits. This review identifies six potential costs to the host from immune activation. The costs are likely to be broadly applicable to other immune responses in vertebrate species. Five phenotypic costs arise from: (i) increased metabolic activity; (ii) reduced nutrient availability due to anorexia; (iii) altered priorities for nutrient utilization; (iv) change in size and turnover of pools of immune cells and proteins; and (v) immunopathology from inappropriate or excessive immune activation. Subsumed by these costs is the cost of altered efficiency of nutrient use. A sixth cost is the genetic cost which arises from a change in the capacity of offspring to express production and life-history traits following selection for parasite resistance. The sensitivity of immune responses to the phenotypic status of the host, and the role the immune system shares with the neuroendocrine system in controlling use of resources underpin the importance of immunocompetence to the life-history of the host.