title: Theoretical analysis of the evolution of immune memory.
abstract: BACKGROUND
  The ability of an immune system to remember pathogens improves the chance of the host to survive a second exposure to the same pathogen. This immunological memory has evolved in response to the pathogen environment of the hosts. In vertebrates, the memory of previous infection is physiologically accomplished by the development of memory T and B cells. Many questions concerning the generation and maintenance of immunological memory are still debated. Is there a limit to how many memory cells a host can generate and maintain? If there is a limit, how should new cells be incorporated into a filled memory compartment? And how many different pathogens should the immune system remember?
  
  
  RESULTS
  In this study, we examine how memory traits evolve as a response to different pathogen environments using an individual-based model. We find that even without a cost related to the maintenance of a memory pool, the positive effect of bigger memory pool sizes saturates. The optimal diversity of a limited memory pool is determined by the probability of re-infection, rather than by the prevalence of a pathogen in the environment, or the frequency of exposure.
  
  
  CONCLUSIONS
  Relating immune memory traits to the pathogen environment of the hosts, our population biological framework sheds light on the evolutionary determinants of immune memory.
authors.affiliation: Integrative Biology, ETH Zurich, Switzerland. frederik.graw@env.ethz.ch
authors.forename: Frederik
authors.initials: F
authors.lastname: Graw
authors.forename: Carsten
authors.initials: C
authors.lastname: Magnus
authors.forename: Roland R
authors.initials: RR
authors.lastname: Regoes
mesh_terms: D005075:Biological Evolution; D003198:Computer Simulation; D056084:Genetic Fitness; D005838:Genotype; D007156:Immunologic Memory / Q000235:genetics*; D018448:Models, Immunological