Optimal virulence

Optimal virulence is a concept relating to the ecology of hosts and parasites. One definition of virulence is the host's parasite-induced loss of fitness. The parasite's fitness is determined by its success in transmitting offspring to other hosts. For about 100 years, the consensus was that virulence decreased and parasitic relationships evolved toward symbiosis. This was even called the law of declining virulence despite being a hypothesis, not even a theory. It has been challenged since the 1980s and has been disproved. A pathogen that is too restrained will lose out in competition to a more aggressive strain that diverts more host resources to its own reproduction. However, the host, being the parasite's resource and habitat in a way, suffers from this higher virulence. This might induce faster host death, and act against the parasite's fitness by reducing probability to encounter another host (killing the host too fast to allow for transmission). Thus, there is a natural force providing pressure on the parasite to "self-limit" virulence. The idea is, then, that there exists an equilibrium point of virulence, where parasite's fitness is highest. Any movement on the virulence axis, towards higher or lower virulence, will result in lower fitness for the parasite, and thus will be selected against. Paul W. Ewald has explored the relationship between virulence and mode of transmission. He came to the conclusion that virulence tends to remain especially high in waterborne and vector-borne infections, such as cholera and dengue. Cholera is spread through sewage and dengue through mosquitos. In the case of respiratory infections, the pathogen depends on an ambulatory host to survive. It must spare the host long enough to find a new host. Water- or vector-borne transmission circumvents the need for a mobile host. Ewald is convinced that the crowding of field hospitals and trench warfare provided an easy route to transmission that evolved the virulence of the 1918 influenza pandemic.