Studies on the ecology of influenza viruses have led to the hypothesis that all mammalian influenza viruses originate from a reservoir of viruses in aquatic birds, particularly ducks. In wild birds, the viruses are spread by fecal–oral transmission through the water supply. Initial transmission of avian influenza viruses to mammals, including pigs, horses, and humans, and to domestic birds, including chickens and turkeys, probably also occurs by fecal contamination of water. Another method of transfer is by feeding pigs untreated garbage or the carcasses of dead birds. After transmission to humans or other mammals, the method of spread of influenza is mainly respiratory.

Influenza represents one of the major success stories for the World Health Organization (WHO). To cope with the variability of influenza, WHO maintains a network of more than 100 laboratories that constantly survey influenza viruses, and this information is then analyzed in four reference centers. Based on these efforts, WHO makes annual recommendations for those virus strains to be included in the current vaccine in order to stay abreast of genetic drift.

The less well resolved problem of influenza is the pandemics, which occur at irregular intervals and are to date unpredictable. In the past century, three viral subtypes have caused pandemics in humans: the Spanish flu of 1918–19, which was caused by the H1N1 subtype; the Asian flu of 1957, which was caused by the H2N2 subtype; and the Hong Kong flu of 1968, which was caused by the H3N2 subtype. The H1N1 and H3N2 subtypes also have caused disease outbreaks in pigs, and the H3N8 and H7N7 subtypes have caused outbreaks in horses.

The Role of Swine in the Emergence of New Influenza Viruses

Generally, influenza viruses are host specific, and viruses from one host rarely establish stable lineages in another host species. Although whole viruses may rarely transmit, gene segments can cross the species barrier through the process of genetic reassortment. Pigs have been postulated to play an important role in the process of genetic reassortment by acting as the “mixing vessel” for such events. Pigs, unlike humans, seem to be readily infected by avian viruses, and most, if not all, avian HA subtypes are capable of replicating in swine. Researchers have proposed a molecular mechanism for the susceptibility of swine to avian virus infection. Viral receptors called sialyloligosaccharides, which are present on the pig tracheal cells, possess the ability to bind to both types of viruses, with human viruses preferentially binding in one location and manner and avian viruses preferentially binding in another location and manner. Thus, pig tracheal cells can be infected not only by human influenza viruses but also by avian viruses. However, the direct chicken-to-human transmission of H5N1 vi-



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