Host Species Barriers to Influenza Virus Infections
Thijs Kuiken1*,
Edward C. Holmes2,
John McCauley3,
Guus F. Rimmelzwaan1,
Catherine S. Williams2 and
Bryan T. Grenfell2,4
1 Department of Virology, Erasmus Medical Center, 3015 GE Rotterdam, Netherlands.
2 Center for Infectious Disease Dynamics, Department of Biology, Pennsylvania State University, University Park, PA 16802, USA.
3 Institute for Animal Health, Compton Laboratory, Compton, Newbury, Berkshire RG20 7NN, UK.
4 Fogarty International Center, National Institutes of Health, Bethesda, MD 20892, USA.
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Fig. 1. Schematic illustrating phases in overcoming species barriers. (A) Interspecific host-host contact must allow transmission of virus from donor species to recipient species. (B) Virus-host interactions within an individual of recipient species affect the likelihood of the virus replicating and being shed sufficiently to infect another individual of recipient species. (C) Intraspecific host-host contact in recipient species must allow viral spread (R0 > 1) in the presence of any preexisting immunity. Superspreader events (red asterisk) early in the transmission chain can help this process. (D) The pathogen must persist in the recipient species population even during epidemic troughs (after most susceptible individuals have had the disease) so that subsequent epidemics can be seeded: If few susceptibles are left, the virus may (stochastically) go extinct in epidemic troughs. Viral variation and evolution can aid invasion and persistence, particularly by affecting host-virus interactions.
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Fig. 2. Evolutionary models for the cross-species transmission of pathogens. (A) The donor and recipient species represent two distinct fitness peaks separated by a steep fitness valley. Multiple adaptive mutations (solid circles) are therefore required for the virus to successfully establish onward transmission in the recipient host species. (B) The donor and recipient species are separated by a far shallower fitness valley. This facilitates successful cross-species transmission because only a small number of advantageous mutations are required. (C) When multiple mutations are required for a virus to adapt to a new host, these may evolve progressively in the recipient species. However, this necessarily requires some onward transmission. (D) It is also possible that many of the mutations required to allow adaptation to a new host species were preexisting in the donor species and transmitted simultaneously. This will accelerate successful viral emergence.
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