Influenza is a highly contagious, acute illness which has afflicted humans
and animals since ancient times. Influenza viruses are part of the Orthomyx
oviridae family and are grouped into types A, B and C according to antigeni
c characteristics of the core proteins. Influenza A viruses infect a large
variety of animal species, including humans, pigs, horses, sea mammals and
birds, occasionally producing devastating pandemics in humans, such as in 1
918, when over twenty million deaths occurred world-wide. The two surface g
lycoproteins of the virus, haemagglutinin (HA) and neuraminidase (NA), are
the most important antigens for inducing protective immunity in the host an
d therefore show the greatest variation. For influenza A viruses, fifteen a
ntigenically distinct HA subtypes and nine NA subtypes are recognised at pr
esent; a virus possesses one HA and one NA subtype, apparently in any combi
nation. Although viruses of relatively few subtype combinations have been i
solated from mammalian species, all subtypes, in most combinations, have be
en isolated from birds. In the 20th Century, the sudden emergence of antige
nically different strains in humans, termed antigenic shift, has occurred o
n four occasions, as follows, in 1918 (H1N1), 1957 (H2N2), 1968 (H3N2) and
1977 (H1N1), each resulting in a pandemic. Frequent epidemics have occurred
between the pandemics as a result of gradual antigenic change in the preva
lent virus, termed antigenic drift. Currently, epidemics occur throughout t
he world in the human population due to infection with influenza A Viruses
of subtypes H1N1 and H3N2 or with influenza B virus. The impact of these ep
idemics is most effectively measured by monitoring excess mortality due to
pneumonia and influenza. Phylogenetic studies suggest that aquatic birds co
uld be the source of all influenza A viruses in other species. Human pandem
ic strains are thought to have emerged through one of the following three m
echanisms:
genetic reassortment (occurring as a result of the segmented genome of the
virus) of avian and human influenza A viruses infecting the same host
direct transfer of whole virus from another species
the re-emergence of a virus which may have caused an epidemic many years ea
rlier.
Since 1996, the viruses H7N7, H5N1 and H9N2 have been transmitted from bird
s to humans but have apparently failed to spread in the human population. S
uch incidents are rare, but transmission between humans and other animals h
as also been demonstrated. This has led to the suggestion that the proposed
reassortment of human and avian viruses occurs in an intermediate animal w
ith subsequent transference to the human population. Pigs have been conside
red the leading contender for the role of intermediary because these animal
s may serve as hosts for productive infections of both avian and human viru
ses and, in addition, the evidence strongly suggests that pigs have been in
volved in interspecies transmission of influenza viruses, particularly the
spread of H1N1 viruses to humans. Global surveillance of influenza is maint
ained by a network of laboratories sponsored by the World Health Organizati
on. The main control measure for influenza in human populations is immunopr
ophylaxis, aimed at the epidemics occurring between pandemics.