Influenza Importance

Influenza viruses are highly variable RNA viruses that can affect birds and
mammals including humans. There are currently three species of these viruses,
Flu, Grippe, Avian Influenza, designated influenza A, B and C. A new influenza C-related virus recently detected in
Grippe Aviaire, Fowl Plague, livestock has been proposed as “influenza D.”1-6
Swine Influenza, Hog Flu, Influenza A viruses are widespread and diverse in wild aquatic birds, which are
Pig Flu, Equine Influenza, thought to be their natural hosts. Poultry are readily infected, and a limited number of
Canine Influenza viruses have adapted to circulate in people, pigs, horses and dogs. In the mammals to
which they are adapted, influenza A viruses usually cause respiratory illnesses with
Last Full Review: February 2016 high morbidity but low mortality rates.7-29 More severe or fatal cases tend to occur
mainly in conjunction with other diseases, debilitation or immunosuppression, as well
as during infancy, pregnancy or old age; however, the risk of severe illness in healthy
Author: humans can increase significantly during pandemics.7,9,11,12,14,20,30-47 Two types of
Anna Rovid Spickler, DVM, PhD influenza viruses are maintained in birds. The majority of these viruses are known as
low pathogenic avian influenza (LPAI) viruses. They usually infect birds
asymptomatically or cause relatively mild clinical signs, unless the disease is
exacerbated by factors such as co-infections with other pathogens.7,46,48-56 However,
some LPAI viruses can mutate to become highly pathogenic avian influenza (HPAI)
viruses, which cause devastating outbreaks of systemic disease in chickens and
turkeys, with morbidity and mortality rates as high as 90-100%.50-52
Although influenza A viruses are host-adapted, they may occasionally infect
other species, and on rare occasions, a virus will change enough to circulate in a new
host. Influenza is now known to be an uncommon cause of respiratory illness in
species not previously considered susceptible, such as cats, ranched mink and various
captive wild mammals, which have been infected by viruses from humans, pigs, birds
and other species.57-77 Dogs have been sporadically affected by viruses from other
species, but were not known to maintain any influenza viruses until the last 20 years,
when two viruses, one from horses and another from birds, began to circulate in some
canine populations.27,63,66,78-90 New viruses have also emerged in pigs, especially in
North America, where swine influenza viruses have become very diverse. 40,91-95 Some
of the North American viruses have affected hundreds of people exposed to pigs at
fairs.96-99 Many influenza A virus infections in novel mammalian hosts have been
mild, but some viruses can cause life-threatening illnesses.53,57-65,97-122 Two
particularly virulent viruses affecting people are the Asian lineage of H5N1 HPAI
viruses and an H7N9 LPAI virus that has caused serious outbreaks in China. Avian
influenza viruses also caused or contributed to at least three past pandemics in
humans, 7,8,14,123-126 while the 2009-2010 pandemic resulted from the acquisition of a
virus from pigs.125,127,128
Influenza B and C viruses primarily affect humans, and seem to be less likely to
cross species barriers.7,11,12,31,129-141 However, infections have occasionally been
reported in animals, and some species may be able to propagate these viruses short-
term, and possibly even for longer periods. 7,16,142-153;154 cited in 146; 155 cited in 156 Little is
known yet about influenza D, but these viruses are thought to circulate in cattle, and
have also been isolated from pigs.1-6 If influenza D viruses are capable of infecting
humans, such infections seem to be rare.1,157
Etiology
Influenza viruses belong to the genera influenzavirus A, influenzavirus B and
influenzavirus C in the family Orthomyxoviridae.158 All the members of each genus
belong to the species influenza A virus, influenza B virus or influenza C virus,
respectively.158 These viruses are also called type A, type B and type C influenza
viruses. A newly recognized livestock influenza virus, originally thought to be an
influenza C virus, might represent a fourth genus, influenza D. 2,3
Influenza A viruses
Influenza A viruses are the most widely distributed influenza viruses in birds and
mammals. These viruses contain two highly variable surface antigens, the
hemagglutinin (HA) and neuraminidase (NA) proteins, which are used to classify

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 Influenza
them into subtypes. Currently, 18 hemagglutinins (H1 to be sufficient for the novel virus to completely evade the
H18) and 11 neuraminidases (N1 to N11) are existing immunity in its host species. Antigenic shifts can
known.50,51,53,159-161 A virus that contained H1 and N2 also occur if one species acquires an influenza virus
would have the subtype H1N2. As of 2016, H1 through “whole” from another, or if a virus disappears for a time
H16 and N1 through N9 have been found in birds; and is maintained in another host species, then re-emerges
H17N10 and H18 N11 viruses have only been detected in in the original host.7,14 For example, human influenza A
bats; and other mammals maintain a limited subset of the viruses can enter and circulate in swine populations, and
subtypes found in birds.15,55,160,161 The HA, and to a lesser could later be re-acquired by humans. In addition to major
extent the NA, are major targets for the immune response, antigenic shifts, reassortment can result in smaller
and there is ordinarily little or no cross-protection between changes, such as the acquisition of a slightly different HA
different HA types or between different NA types. 9,162-170 or NA from a similar virus circulating in the same species,
Considerable variation also exists within each HA or NA or a different internal protein.
type, and two viruses that share a subtype may be only Antigenic drift and shifts result in the periodic
distantly related. emergence of viruses with new or altered HA and NA
The names of influenza A viruses reflect the hosts to proteins. By evading the immune response, these viruses
which they are adapted. Influenza A viruses are currently can cause influenza epidemics and pandemics.
maintained in birds (avian influenza viruses), pigs (swine Acquisition and loss of influenza viruses in
influenza viruses), horses (equine influenza viruses), dogs a species
(canine influenza viruses) and humans. The viruses Each influenza A virus is adapted to circulate in a
adapted to people are called human influenza A viruses, to particular host; however, viruses can occasionally infect
distinguish them from influenza B and C viruses, which other species. In most cases, the virus cannot be
are also maintained in human populations. Together, the transmitted efficiently in the novel host, and soon
influenza A, B and C viruses circulating in people are disappears.7,8,15,27,46,53,65,74,75,81,89,171-173 On rare occasions,
called ‘seasonal influenza viruses.’ however, a virus continues to circulate. Complex
Strains of influenza viruses are described by their molecular adaptations, which are still not well understood,
type, host, place of first isolation, strain number (if any), are likely to be required for a successful species jump.174
year of isolation and subtype.7,15 For example, the The viral surface proteins (HA and NA) and internal
prototype strain of the H7N7 subtype of equine influenza proteins both seem to be involved in host preferences.
virus, first isolated in Czechoslovakia in 1956, is Viruses generally undergo a period of adaptation after the
A/eq/Prague/56 (H7N7). For human strains, the host is transfer, during which time they become more efficient at
omitted. When an influenza virus lineage has circulated replicating in the new host.
for a time in a population, numerous variants may develop. In some cases, whole viruses have jumped
These variants are sometimes classified into clades and successfully to new species.27,79 At other times, the newly
subclades (e.g., clade 2.2 of the Asian lineage H5N1 HPAI acquired virus reassorted with a virus already adapted to
virus). its new host.8,14 Reassortment can occur either in the new
Antigenic shift and drift in influenza A viruses host or in an intermediate host, which then transmits the
Influenza A viruses are very diverse and change virus further.8,14,53 For example, an avian influenza virus
frequently as the result of two processes, mutation and could reassort with a human influenza virus in a pig, then
genetic reassortment. Mutations cause gradual changes in be transferred to humans. Acquisition of new influenza
the HA and NA proteins of the virus, a process called viruses is more likely when different species are kept in
‘antigenic drift.’15 Once these proteins have changed close proximity.7,14,108
sufficiently, immune responses against the former HA and As well as appearing in new host populations,
NA may no longer be protective. influenza A viruses can disappear from hosts where they
Genetic reassortment can cause more rapid changes. previously circulated. Some viruses have vanished from
The influenza A genome consists of 8 individual gene human, equine and swine populations after circulating for
segments,169,170 and when two different viruses infect the years or even decades.7,13,93-95 For unknown reasons, the
same cell, gene segments from both viruses may be establishment of a new influenza virus in a species
packaged into a single, novel virion.8 This can occur sometimes leads to the disappearance of an older viral
whenever two influenza viruses replicate in the same cell, lineage.39
whether the viruses are adapted to the same host species Avian influenza viruses
(e.g., two different human influenza viruses) or originally Avian influenza viruses are extremely diverse,
came from different hosts, such as an avian influenza virus especially in wild birds, and can contain any of the HA
and a swine influenza virus. and NA proteins other than H17, H18, N10 or N11.
An important aspect of reassortment is that it can Whether all HA and NA combinations can occur in nature
generate viruses containing either a new HA, a new NA, or is uncertain, but more than 100 subtypes of avian influenza
both. Such abrupt changes, called ‘antigenic shifts,’ may viruses have been detected.39 Some hemagglutinins, such

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 Influenza
as H14 and H15, seem to be uncommon, or perhaps are (e.g., a highly divergent lineage first identified in
maintained in wild bird species or locations that are not Argentina).197-199 The viruses in New Zealand and
usually sampled.55 Australia might be geographically isolated to some extent,
although there is also evidence of mixing with viruses
LPAI and HPAI viruses
from other areas.200-202
The viruses maintained in birds are classified as either
low pathogenic (also called low pathogenicity) avian Important virus lineages circulating among poultry:
influenza viruses or highly pathogenic (high Asian lineage H5 HPAI viruses, and H7N9 and H9N2
pathogenicity) avian influenza viruses. A virus is defined LPAI viruses
as HPAI by its ability to cause severe disease in Many different LPAI and HPAI viruses, belonging
intravenously inoculated young chickens in the laboratory, to multiple subtypes, can infect poultry, but three
or by its possession of certain genetic features that have lineages of H5, H7N9 and H9N2 viruses are currently of
been associated with high virulence in HPAI viruses (i.e., particular concern.
the sequence at the HA cleavage site).50,51 HPAI viruses Asian lineage H5 HPAI viruses
almost always cause severe disease when they infect The A⁄goose⁄Guangdong⁄1996 lineage (‘Asian
chickens and turkeys in the field, while LPAI infections lineage’) of H5N1 HPAI viruses first emerged among
are generally much milder. poultry in China in the late 1990s, and has become
With rare exceptions, the HPAI viruses found in widespread and diverse.53,203,204 These H5N1 viruses have
nature have always contained the H5 or H7 evolved into multiple genotypes, clades and subclades, and
hemagglutinin.175-178 Two exceptions were H10 viruses new variants are continuing to emerge as they
that technically fit the HPAI definition if they were circulate.53,204-209 Some clades or subclades differ in their
injected directly into the bloodstream of chickens, but virulence for mammals and/or birds.110,121,208 The primary
caused only mild illness in birds infected by the reasons for concern, in addition to the severe outbreaks
respiratory route.177 Another H10 virus that fit the HPAI HPAI viruses cause in poultry, are the serious illnesses
definition affected the kidneys and had a high mortality H5N1 viruses cause in humans, the wide variety of
rate even in intranasally inoculated chickens.179 In the mammalian species they can infect, and the periodic (and
laboratory, the insertion of genetic sequences from HPAI unusual) detection of these viruses in wild birds, including
viruses into non-H7, non-H5 viruses has created some migratory waterfowl.57,58,60-65,100-121,210,211
viruses that were pathogenic only after intravenous Asian lineage H5N1 viruses have reassorted with
inoculation, and other viruses (containing H2, H4, H8 or other avian influenza viruses, and several new subtypes
H14) that were highly virulent after both intravenous and belonging to this lineage (e.g., H5N2, H5N5, H5N6 and
intranasal inoculation.180 Recently, an H4N2 virus with a H5N8) have been found in poultry in Asia.212-218 Asian
genetic signature characteristic of HPAI viruses was lineage HPAI H5N8 viruses became widespread, and
isolated from a flock of naturally infected quail.181 This reached Europe and North America in 2014, most likely in
virus retained the characteristics of an LPAI virus, and was wild birds.219,220,220-222 These viruses have reassorted with
of low virulence in chickens. North American lineage viruses to produce several new
In rare cases, an H5 or H7 virus has a genetic variants, including H5N2 HPAI viruses, which caused
signature that classifies it as an HPAI virus, but causes extensive outbreaks among poultry in the Midwestern
only mild illness in poultry.182,183 These viruses may have U.S.220-228 Some H5 reassortants in Asia have caused
been isolated when they were evolving to become more illnesses in mammals. They include an H5N2 virus
virulent. Their presence triggers the same regulatory isolated from a sick dog, and H5N6 viruses in several
responses as fully virulent HPAI viruses. human cases.66,229-234 As of February 2016, no clinical
Avian influenza virus lineages cases have been linked to H5N8 viruses except in birds.
There are two well-recognized lineages of avian Eurasian lineage H9N2 viruses
influenza viruses, Eurasian and North American.55 As A Eurasian lineage of H9N2 (LPAI) viruses is
implied by the names, Eurasian lineage viruses primarily currently widespread among poultry in parts of Asia and
circulate among birds in Eurasia, and North American the Middle East. These viruses have caused clinical cases
lineage viruses in the Americas. The amount of (mostly mild) in people, and they can infect other
reassortment between these lineages seems to differ mammals.94,95,235-252 They have also become very diverse,
between regions, with very few reassortant viruses and there is evidence of reassortment with Asian lineage
detected in some areas or wild bird populations, but H5 HPAI viruses and other lineages. 236,245,253-257 H9N2
significant reassortment where there is overlap between variants may differ in their ability to replicate in
migratory flyways, such as in Alaska and Iceland. 55,184-196 mammals and/or cause disease. 252,255,256
Limited information from Central and South America
suggests that many or most of the viruses in this region are
closely related to the North American lineage, but H7N9 LPAI viruses
cocirculate with some viruses unique to South America

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 Influenza
A lineage of H7N9 LPAI viruses in China causes little 2009 pandemic H1N1 virus
or no disease in poultry, but has been responsible for A novel H1N1 virus emerged in human populations in
serious outbreaks in humans. 258-264 These viruses seem to 2009 and caused a pandemic.125,127,128,275 This virus is
have arisen from reassortment between H7, N9 and H9N2 thought to be a reassortant between North American H1N2
viruses.259,265 Some evidence suggests that they may have and Eurasian H1N1 swine influenza viruses.125,127,128 Its
circulated subclinically for a long time among poultry HA is most closely related to swine influenza viruses in
before emerging in people.266 H7N9 viruses have North America, and the NA to swine influenza viruses in
diversified considerably since they were first recognized, Eurasia, while the internal proteins originated from two or
and regionally distinct lineages now exist in China. 267 more swine influenza viruses including the North
H7N7 viruses with similar internal H9N2 genes have been American triple reassortant H3N2 viruses (see swine
identified among poultry in China, and might also have the influenza, next section) and a Eurasian virus. Some of the
potential to infect mammals.265 gene segments in these swine viruses originally came from
Human influenza A viruses avian and human influenza viruses.125,128 How humans
acquired pandemic H1N1 virus is not known, but genetic
In people, influenza A viruses tend to form a single
analyses suggest that it was probably transmitted to people
global population.248 H1N1, H1N2, N2N2 and H3N2
shortly before the pandemic began, most likely from pigs,
human influenza viruses have been widely distributed at
and that it might have previously circulated among swine
times during the last century, but only H1N1 and H3N2
in an unknown location for years.125,128 After the 2009-
viruses are currently in general circulation.7,30,31,268,269
2010 pandemic, this virus became established as a
Human influenza viruses are under considerable seasonal influenza virus, and it continues to circulate
selection pressure from immunity (via infection or throughout the world. It is currently the predominant
vaccination) in a long-lived species. As a result, the circulating H1N1 virus in people.276 This virus has had
predominant viruses circulating in human populations several names (e.g., swine influenza virus, swine-origin
change constantly, resulting in epidemics and pandemics. influenza virus, novel H1N1) but, at present, the most
Pandemics were most recently reported in 1918, 1957, commonly used name is 2009 pandemic influenza A
1968 and 2009. The 1918 ‘Spanish flu’ pandemic was (H1N1) virus or 2009 pandemic H1N1 virus.
caused by an H1N1 virus whose origins remain
controversial. Some evidence suggests that it was Swine influenza viruses
probably an avian virus that became adapted to At present, diverse viruses of the subtypes H1N1,
humans,8,123 while other studies indicate that it may have H1N2 and H3N2 circulate in swine populations, although
been a reassortant,125,126,270 and it is possible that it other subtypes have transiently infected pigs in limited
adapted to another host, such as a pig, before becoming locations.8,14,40,46,91,93-95,248,277-282 Different sets of viruses
established in people. 125,271 H1N1 viruses gradually circulate on each continent, and sometimes in different
changed as they circulated in the human population, then countries or regions within a continent. 8,14,40,46,93-95,248,282-
284
apparently disappeared in 1957 when an H2N2 virus
emerged.8,14,124,272 The next two pandemics seem to have North America
been caused by reassortment between avian and human
The first influenza virus recognized in pigs was an
influenza viruses. 8 The 1957 H2N2 (‘Asian flu’) virus
H1N1 virus known as the ‘classical’ swine influenza
consisted of the HA, NA and an internal protein from an
virus.285-287 Pigs are thought to have acquired this virus
avian influenza virus, and five other proteins from a
during the 1918 Spanish flu pandemic, possibly from
human H1N1 strain. 8,14 These H2N2 viruses circulated in
infected people.7,8,14,46,270,286-289 H1N1 viruses circulated in
people between 1957 and 1968. 7,269 The H3N2 ‘Hong
both species after this time, but diverged genetically in the
Kong flu’ virus, which appeared in 1968, had two new
two host populations.272,290
proteins from avian viruses - the new HA and an internal
protein - but kept the NA and remaining proteins from The classical H1N1 swine influenza virus was the
the H2N2 virus. 8,14 H1N1 viruses re-emerged into human major virus among swine populations in North America
populations in 1977, and then co-circulated with the for approximately 70 years.40 Low levels of H3 viruses
H3N2 viruses.7,124 (While this event is also technically a from humans were also found during this period, and some
pandemic, these viruses were not new, but descendants of of these viruses circulated locally for at least a short time;
the H1N1 viruses that first entered human populations in however, they did not become established as stable,
1918.) H1N2 viruses have not caused a pandemic, but widespread lineages.40,291 Triple reassortant H3N2 viruses
viruses with this subtype been found at times in limited first emerged in North American pigs in the late 1990s,
locations, and one H1N2 virus (which probably resulted mainly in the U.S. Midwest, and spread to other
from genetic reassortment between H3N2 and H1N1 regions.8,74,108,277,292-295 These viruses contain HA and NA
viruses) circulated globally between 2001 and derived from human influenza viruses, and internal
2003.30,269,273,274 proteins from the classical swine influenza virus, an avian
influenza virus and a human influenza virus.294 The
particular combination of internal genes carried by these

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 Influenza
viruses is known as the triple reassortant internal gene available.94 A long-term analysis conducted in Hong Kong
(TRIG) cassette. Viruses that contain TRIG seem to be abattoirs, where most of the pigs originate from China,
prone to increased antigenic drift.92 They also seem to suggests that swine influenza viruses reassort frequently,
readily acquire new HA and NA genes; there are now but only a few of these viruses persist, and that the
additional TRIG-containing swine influenza viruses with population of viruses gradually changes.248 This is also
various combinations of H1, H3, N1 or N2 from additional likely to be true of other regions and continents.
human influenza viruses, and/or H1 and N1 from the
Other locations
classical swine influenza virus.40,91,92,275,296,297 Some North
American herds have been infected with the 2009 pandemic At present, there is little information about swine
H1N1 virus from humans, which has reassorted with other influenza viruses in some parts of the world. H3N2 and
viruses circulating in pigs.75,298-303 As a result of all these H1N1 viruses are known to circulate in Latin America,
factors, North American H1N1, H1N2 and H3N2 swine but limited information is available about their genetic
influenza viruses have become very diverse, and are composition.319 A recent study found that viruses in
continuing to change.40,91,92,296,297,303,304 Other influenza northwestern Mexico included H3N2 viruses similar to
variants and subtypes, such as H2N3 and H3N1 viruses, those in the U.S., and H1N1 viruses that had 89%
have been found occasionally in North American herds, but nucleotide identity with U.S. viruses. 320 One H3N2 virus
do not seem to have become established in swine isolated from sick pigs in Argentina was of wholly
populations.277-281 human origin, although it was highly transmissible in
pigs.319 Whether this was a limited outbreak, or the virus
Europe circulates there in swine populations is not known.
The classical H1N1 swine influenza virus was found Pandemic H1N1 and/or its reassortants with human-like
sporadically in Europe in the past (although records of its H1N1 swine influenza viruses, as well as a reassortant
isolation and times of circulation are scarce), but a virus of human-like H1N2 influenza virus, have been detected in
wholly avian origins has been the major H1N1 swine Argentina and Brazil. 321-324 H1 viruses have been
influenza virus since the 1970s.8,14,39,46,93,94,282-284,305-308 The documented in Africa, and some recent studies report the
2009 pandemic H1N1 virus and its reassortants are also presence of the 2009 pandemic H1N1 virus there in
detected frequently, and currently seem to be maintained pigs.325-327
in swine populations in some areas.39,93,282-284,309 Various
Equine influenza viruses
human-origin H3N2 viruses have circulated at times in
pigs, but a reassortant that contains human-origin H3 and Equine influenza viruses seem to change more slowly
N2 with internal gene segments from the avian-origin than human influenza A viruses or swine influenza
H1N1 virus has now become the dominant H3N2 virus in viruses.13,15,328,329 Two subtypes of influenza viruses
much of Europe.14,39,93,282-284,305 Several H1N2 viruses circulated widely in equine populations during the last
(including pandemic H1N1 reassortants) also occur, either century, H7N7 (equine virus 1) and H3N8 (equine virus
transiently or long-term.14,39,93,282-284,305,309,310 Additional 2).7,13,15 H7N7 equine influenza viruses were last isolated
subtypes (e.g., H3N1 viruses) have been found in 1979, and most authors believe they are likely to be
occasionally.39,93,311 One unique variant was an H1N7 extinct, although there have been a few anecdotal reports
virus, which was apparently a reassortant between swine or serological studies suggesting that they might persist in
and equine influenza viruses.39,93 some areas where surveillance is limited.7,13,19,328,330,331;332
cited in 333

Asia In the 1980s, equine H3N8 viruses diverged into

Information about swine influenza viruses in Asia is distinct Eurasian and American evolutionary lineages.334-
limited, especially for some regions, but H1N1, H3N2 and 336
The American lineage divided further into 3
H1N2 viruses are known to circulate. Various North sublineages: the classical American (or Kentucky) lineage,
American and European lineage viruses belonging to these the Florida sublineage and the South American
three subtypes have been reported, as well as reassortants sublineage.329,335,336 The Florida sublineage became
between North American and Eurasian lineages, and widespread, and has diverged into 2 clades.329,334-336 The
viruses unique to Asia.46,65,94,95,248,312-316 Some of the Eurasian lineage is now uncommon (some reports indicate
circulating viruses contain the North American TRIG that it was last isolated in 2005), and the classical
cassette. Some viruses have infected Asian pigs only American lineage is found occasionally in some
transiently, and different swine influenza viruses may areas.329,334-336
predominate in different regions.94,95 The pandemic H1N1 A novel H3N8 virus (A/eq/Jilin/89) caused two
virus and its reassortants have also been found, and novel outbreaks in China in the late 1980s/ early 1990s, with
subtypes (e.g., H3N1 viruses) have been isolated serological evidence indicating continued exposure of
occasionally.94,95,317,318 horse populations for a few years longer.13,19,65 This virus
Whether a virus is circulating in pigs or represents a appeared to be of avian origin. It did not persist long-term
one-time event can sometimes be difficult to determine and is not known to have spread outside China.13
without long-term surveillance, which is not always

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 Influenza
Canine influenza viruses viruses, was recently isolated from pigs and cattle.1,2
The first canine influenza virus to be recognized was Initially, this virus was thought to be a new subtype of
an H3N8 (Florida sublineage) virus acquired from horses influenza C, but subsequent studies suggest that it might
in North America in the late 1990s or early 2000s. 27,79,80 represent a new genus of influenza viruses.2-4 The name
This virus has diverged genetically from equine influenza “influenza D” has been proposed,2,3 although it is not yet
viruses, and adapted to circulate in dogs.27,79,337-339 It may formally accepted by the ICTV.158 At least two lineages of
now be evolving into two lineages.340 influenza D viruses have been found in North American
An H3N2 canine influenza virus, with gene segments cattle, and these lineages can reassort.355 Influenza D
entirely of avian origin, became established in some Asian viruses found in China, Italy and France were closely
countries in the mid-2000s.76,81-88,341 These viruses have related to viruses from North America.4-6
diversified and may be evolving separately in different
Species Affected
countries.342 They have also reassorted with other viruses.
An H3N1 virus, which appears to be a reassortant between Avian influenza viruses
a canine H3N2 virus and the human pandemic H1N1
Wild birds
virus, was isolated from one dog with respiratory signs in
Korea.343 There is also evidence of occasional Birds are thought to be the natural reservoir hosts from
reassortment with avian influenza viruses from poultry which all influenza A viruses originated.7,46,49,51,54,55,108 The
(e.g., H9N2, H5N1).341,342,344 vast majority of LPAI viruses are maintained in
asymptomatic wild birds, particularly species found in
Bat influenza viruses wetlands and other aquatic habitats.7,46,49,51,54-56,159,356
Two new subtypes of influenza viruses were identified Infections are especially common in some members of the
recently in South American bats, using genetic order Anseriformes (waterfowl, such as ducks, geese and
techniques.160,161 These viruses have distinct swans) and two families within the order Charadriiformes,
hemagglutinins, which have provisionally been designated the Laridae (e.g., gulls) and Scolopacidae
H17 and H18, unique neuraminidases, and distinctive (shorebirds).7,14,46,49,51,55,56,159,188,193,196,201,356,357,357-361 Aquatic
internal genes.160,161 They cannot be isolated in the cell lines species belonging to other orders occasionally have high
and chicken embryos used to grow other influenza A infection rates, and might also be involved in the
viruses. The inability to recover infectious virus has made it epidemiology of this disease.56,362,363; 364 cited in 363 While LPAI
difficult to study bat influenza viruses; however, various viruses can infect birds that live on land (terrestrial birds),
analyses suggest that they are unusual and differ from other most studies have reported low infection rates, suggesting
influenza A viruses in multiple ways.160,345-348 that these birds are not important reservoir hosts in most
environments.56,356,365-374 Higher rates are occasionally
Influenza B viruses
reported in individual species, and in a study from Vietnam,
Influenza B viruses are categorized into lineages (and viruses were particularly common in some terrestrial birds
strains) based on the HA, but not into subtypes. Currently, that forage in flocks.367,373 Recently, influenza virus RNA
the two important lineages in people are represented by was found in an unusually high percentage of passerine
B/Victoria/2/87 and B/Yamagata/16/88.349 Both lineages birds in Central and West Africa.375
are currently widespread and co-circulate, although one
The most common influenza subtypes in wild birds
lineage may predominate in an area in any given year. 349-
351 may differ between species and regions, and can change
Recombination can occur within, and occasionally
over time.55,359,360,363,376-378 Some birds may maintain
between, the two lineages; and influenza B viruses can
viruses long-term, while others might be spillover hosts.
undergo infrequent antigenic shifts.350,352,353 Antigenic drift
Migrating birds, which can fly long distances, may
also occurs, although it generally happens more slowly
exchange viruses with other populations at staging,
than in influenza A viruses.7,268,353
stopover or wintering sites.55 A few avian influenza
Influenza C viruses subtypes seem to have a limited host range, and may rarely
Influenza C viruses have one surface glycoprotein, the (or never) be transferred to poultry.55,184,188,360,379-385
hemagglutinin-esterase-fusion protein, rather than HA and HPAI viruses are not usually found in wild birds,
NA proteins. They are antigenically more stable than although they may be isolated transiently near outbreaks in
influenza A viruses, and accumulate fewer changes over poultry.369 Exceptions include the Asian lineage H5N1
time.130,354 At least six lineages (Taylor/1233/47-, Sao viruses and some of their reassortants (e.g., H5N8 viruses),
Paolo/378/82-, Kanagawa/1/76-, Aichi/1/81-, which have been found repeatedly in wild birds, an H5N3
Yamagata/26/81- and Mississippi/80-related lineages) virus isolated from an outbreak among terns in the 1960s,
have been identified.354 Reassortment can occur between an H7N1 virus that was isolated from a sick wild siskin,
different strains or lineages.130,131,157,354 Carduelis spinus, and an H5N2 virus found in a few
asymptomatic wild ducks and geese in
Influenza D (Influenza C-related livestock virus) Africa.62,101,108,109,119-121,219,221,222,224-226,228,386,387,387-404
A new livestock-associated influenza virus, which
shares 50% amino acid identity with human influenza C

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 Influenza
Domesticated birds replicated inefficiently in experimentally infected dogs,
Poultry and game birds, including gallinaceous birds but cats were more likely to become infected in this
and domesticated waterfowl, are readily infected by LPAI study.435 Initial laboratory experiments in ferrets and mice
and HPAI viruses.51,109,119-121,165,395,405-419 When LPAI suggested that these H5N8 viruses may be less pathogenic
viruses from wild birds are transferred to poultry, the for mammals than some H5N1 isolates.435-437 Asian
viruses may circulate inefficiently and die out; become lineage H5N6 viruses have been isolated from apparently
adapted to the new host and continue to circulate as LPAI healthy pigs.438
viruses; or if they contain H5 or H7, they may evolve into Host range of Eurasian H9N2 (LPAI) avian
HPAI viruses.53,54,175 Infections also occur in other influenza viruses
domesticated birds, although they do not appear to be very H9N2 viruses have been detected in poultry and some
common in either cage birds or wild birds including terrestrial species.245,439,440 They have
pigeons.48,101,109,119,370,390,396,420-424 Once a virus has adapted also been found in pigs, dogs and farmed
to poultry, it rarely re-establishes itself in wild birds.175 mink,94,95,236,246,247,249,250,252,441,442 and serological evidence of
Host range of the Asian lineage H5N1 HPAI avian infection has been reported in cats, captive nonhuman
influenza viruses and reassortants including H5N8 primates (performing macaques) and wild plateau
Asian lineage H5N1 HPAI viruses have an unusually pikas.251,443,444 Experimental infections have been
wide host range. These viruses can infect domesticated and established in mink and pika.441,444 Dogs and cats could also
wild birds belonging to many different orders.62,101,108,109,119- be infected experimentally with some isolates, although
121,391-400
They can be highly virulent in some birds, but virus replication was limited in some studies.445-447
some infections are subclinical.101,396,399,403,425 Likewise, Host range of the zoonotic H7N9 avian influenza
Asian lineage H5N8 viruses and/or their reassortants have viruses
been detected in various domesticated and wild birds The zoonotic H7N9 LPAI viruses in China have
including sick, dead and apparently healthy waterfowl, and mainly been found in poultry, although live viruses and/or
sick or dead birds in other orders including viral nucleic acids were also detected in a few wild birds
raptors.219,221,224,226,386,387,401,402,404 Wild migratory birds have (e.g., two pigeons, an asymptomatic tree sparrow, and wild
introduced some Asian lineage H5 viruses (e.g., H5N1 and waterfowl).264,448,449 Based on experimental infections,
H5N8 viruses) to uninfected regions, but whether they can chickens and quail are most likely to maintain this virus,
maintain these viruses for long periods (or indefinitely), or but several species of ducks, geese, pigeons, parakeets
are repeatedly infected from poultry, is still (Melopsittacus undulates) and various passerine birds
controversial.219,219,220,222,225,228,386,387,391,394,399,426-428 could also be infected.424,450,451 H7N9 viruses are not
Asian lineage H5N1 HPAI viruses can infect many reported to have caused any illnesses in mammals other
species of mammals, and their full host range is probably than humans,264 as of February 2016. Stray dogs living
not yet known. They have been found in pigs, cats, dogs, near live poultry markets did not appear to have been
donkeys, tigers (Panthera tigris), leopards (Panthera infected,246 and a serological survey found little or no
pardus), clouded leopards (Neofelis nebulos), lions evidence of exposure in pigs.452 In experimental studies,
(Panthera leo), Asiatic golden cats (Catopuma isolates from humans could infect miniature pigs, ferrets,
temminckii), stone martens (Mustela foina), raccoon dogs laboratory mice and cynomolgus macaques.451,453-455
(Nyctereutes procyonoides), palm civets (Chrotogale Experimentally infected black rats (Rattus rattus) did not
owstoni), plateau pikas (Ochotona curzoniae) and a wild shed measurable amounts of virus, although they
mink (Mustela vison).53,57-65,100-104,106,107,122,211,246,429,430 seroconverted.451
Antibodies to these viruses were detected in horses and
raccoons.65,104,106,107,122,246,430-433 Experimental infections Other avian influenza viruses in mammals
have been established in housecats, dogs, foxes, pigs, Other avian influenza viruses are reported
ferrets, laboratory rodents, cynomolgus macaques sporadically in mammals. Diverse subtypes (e.g., H4,
(Macaca fascicularis) and rabbits.65,101,105,106,108-118,429,434 H5N2, H6N6, H7, H10N5 and H11N2) have been isolated
Viruses isolated from cats could infect cattle in the occasionally from pigs, especially in Asia.94,95,171,236,452,456-
460
laboratory,118 but serological studies in Egypt suggest that An H10N4 virus was responsible for an epidemic
cattle, buffalo, sheep and goats are not normally infected among farmed mink in Europe,65 and experimental
with H5N1 viruses.431 infections with H3N8, H4N6, H5N3, H7N7, H8N4 and
Some Asian lineage H5 reassortants, such as an H5N2 H11N4 avian influenza viruses have been established in
virus, recovered from a sick dog in China, may cause this species.7,65 Cats have been infected experimentally
illnesses in mammals.66,229,230 This H5N2 virus could be with H1N9, H6N4, and H7N3 LPAI viruses, as well as
transmitted from experimentally infected dogs to dogs, with an H7N7 HPAI virus isolated from a fatal human
chickens and cats.66,229,230 There have been no reports of case.461-463 Some dogs in China has antibodies to H10N8
illnesses caused by Asian lineage H5N8 viruses, as of viruses,464 and surveillance found an H6N1 LPAI virus in
February 2016, although seropositive dogs were detected one dog coinfected with canine distemper virus in
on some infected farms in Asia.435 An H5N8 virus Taiwan.465 Dogs were also susceptible to experimental

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 Influenza
infection with an H6N1 virus.466 Some domesticated coast of North America in 2010, but there was no evidence
guinea pigs in South America had antibodies to H5 of infection in other marine mammals tested.528
influenza viruses.152 Experimental infections have been established in cats,
Few studies have investigated wild animals; however, dogs, ferrets, mice, cynomolgus macaques, turkeys and
antibodies to H4 and H10 viruses were found in raccoons quail.523-526,529-532 Chickens do not seem to be susceptible
in the U.S. (in addition to antibodies to H1 and H3 viruses, to this virus.530,531,533
which could also originate from mammals), and antibodies Swine influenza viruses
to H3N8 viruses, possibly of avian origin, were reported in
Swine influenza viruses mainly affect pigs, but some
raccoons in Japan.467-469 Raccoons could be infected
viruses can also cause disease in turkeys, ferrets and
experimentally with an avian H4N8 LPAI virus, striped
mink.7,15,65,74,75,171,173,278,290,503,534-538 Some swine influenza
skunks (Mephitis mephitis) with H4N6 and H3N8 LPAI
viruses found in turkeys can be transmitted back to pigs,
viruses, and cottontail rabbits with an H4N6 LPAI
but others no longer readily infect this species. 171 Chicken
virus.468,470,471;472 cited in 470 Avian influenza viruses have
flocks infected with swine influenza viruses have not been
been associated with disease outbreaks in seals (see
reported, and chickens do not seem to be very susceptible
Marine Mammals, below).
to these viruses after experimental inoculation.171 One
Laboratory mice (Mus musculus) and ferrets serve as H1N1 swine influenza virus, which was avirulent for both
models for mammalian infections with influenza viruses, poultry and pigs, was isolated from a duck in Hong
including avian influenza viruses. 473-481 Most laboratory Kong,539 and ducks can be infected experimentally.497 Two
mice have a defective gene (Mx1), which increases their H3N2 viruses isolated from pet dogs in China had high
susceptibility to influenza viruses compared to their wild- homology to human-like H3N2 swine influenza viruses
type progenitors. 482-484 However, one recent study found among pigs in the area.540 Experimental infections
suggested that wild Mus musculus mice may also be have been reported in calves.539,541 Antibodies to H3
susceptible to experimental inoculation with certain viruses found recently in cattle might have been caused by
LPAI viruses.482 exposure to swine influenza viruses, but definitive
Human influenza viruses identification of the virus source was not possible.542
Human influenza A viruses mainly cause disease in Equine influenza viruses
people, but nonhuman primates are also susceptible to these Equine influenza viruses mainly affect horses and
viruses, and pet ferrets can become ill.10,153,251,485-488 Most other Equidae (e.g., donkeys, mules and zebras). 7,13,18,543 In
primate studies have been done in captive animals, but there several instances, dogs have been infected by H3N8
was also evidence of infections in pet, performing and wild viruses from nearby horses, without the virus becoming
macaques in Asia.251 Human influenza virus infections are established in canine populations.24,89,90,544,545 One H3N8
reported occasionally in pigs, and human or part-human virus was found during surveillance of healthy Bactrian
viruses can become established in these animals.8,14,39,40,93- camels (Camelus bactrianus),546 another H3N8 virus was
95,248,277,282,291,294,313,314,319,489
Serological and/or virological isolated from sick pigs in China,547 and a reassortant
evidence of infection has been reported sporadically in other between swine and equine influenza viruses (H1N7) was
animals such as dogs, cats, cattle and even birds (including detected in pigs in Europe.39,93 Experimental infections
poultry),7,15,142,151,153,251,277,313,331,490-493 and experimental have been established in dogs, cats, ferrets and
infections have been established in cats, dogs, mink, mink;65,77,478,548,549 however, one group found that an
raccoons and ducks.7,19,468,488,491,493-497 Some domesticated equine H3N8 virus did not replicate well in experimentally
guinea pigs in South America have antibodies to H1 and H3 infected swine.550 Cattle were susceptible to an equine
viruses that might be of human origin.152 Human influenza influenza virus in an older experiment,7,19 but in a recent
A viruses can replicate, to a limited extent, in the nasal study, there was no evidence of infection when they were
epithelium of experimentally infected horses,19 and there exposed to an aerosolized H3N8 virus.542
was some evidence for inapparent infections in horses at the
time of the human ‘Asian flu’ epidemic.498 Canine influenza viruses
The 2009 pandemic H1N1 virus (now a seasonal The H3N8 canine influenza virus has only been
human influenza virus) is often found in domesticated reported in dogs.22,23,25,27,28,79 One group reported finding
pigs,93,95,298-300,309,322,326,499-502 and has been detected in wild viral nucleic acids in two cats, but concluded that the cats
boar.503 It has caused outbreaks in turkeys,283,284,490,504-509 were not infected as they never developed measurable
and a few clinical cases have been reported in pet ferrets, antibody (HI) titers to the virus.551 Although the H3N8
cats and dogs; captive wild species including cheetahs, a canine influenza virus can still infect horses under some
black-footed ferret (Mustela nigripes), an American experimental conditions, its ability to replicate in equids
badger (Taxidea taxus taxus), a Bornean binturong appears to be significantly reduced.338,339 One study
(Arctictis binturong penicillatus), captive giant pandas reported that horses were not infected when kept in close
(Ailuropoda melanoleuca); and possibly wild striped contact with experimentally infected dogs.552 In laboratory
skunks.67-73,510-527 This virus was found in healthy wild studies, this virus was not transmitted readily to chickens,
northern elephant seals (Mirounga angustirostris) off the turkeys or ducks,553 and it did not replicate well in pigs.550

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 Influenza
The H3N2 canine influenza virus has caused clinical Influenza B viruses
cases in dogs and cats.76,81,86,87,554 Antibodies to this virus Virological and/or serological evidence of, influenza B
have been found in both species in Asia, dogs and cats are infections has been reported in seals, pigs, dogs, horses,
susceptible to experimental infection by contact with captive nonhuman primates, guinea pigs and some avian
infected dogs, and cats can transmit the virus to other species, and these viruses have been associated with
cats.76,81-88,246,555-558 Ferrets have also been infected illnesses in ferrets, seals and experimentally infected
experimentally, but seem to be less susceptible, and pheasants7,14,16,65,142,145-153,567;154 cited in 146; 568-570 cited in 150; 155 cited
animal-to-animal transmission only occurred to a limited in 156
One recent study suggested that influenza B viruses
extent.556,559,560 Guinea pigs can also be infected might be more common in pigs when the animals are co-
experimentally; however, attempts to transmit the H3N2 infected with PRRS virus.149 Experimental infections have
canine influenza virus to chickens, ducks, mice and pigs been established in pigs,149,156 horses,571 guinea pigs,572
were unsuccessful.83,560 pheasants, 150 mallard ducks (Anas platyrhynchos) 150 and
Influenza viruses in marine mammals chickens.150 Older studies also described experimental
infections in cats, dogs and nonhuman primates,494,496,573-575
A number of influenza viruses (H3N3, H3N8, H7N7, cited in 150
and one group recovered an influenza B virus from
H4N5, H4N6 and H10N7), which were closely related to
dogs during an outbreak in Japan.151 However, one recent
avian viruses, have been isolated from seals, and human
study found no evidence of productive infection in
pandemic H1N1 virus was found in northern elephant
experimentally infected dogs.493
seals in 2010.7,65,528,561-564 An avian-origin H3N8 virus,
isolated during North American outbreak in 2011, Influenza B viruses are proven to circulate only in
appeared to have adaptations that would increase its human populations, although some evidence suggests
transmissibility in mammals.562 This virus also replicated that either seals or an unknown marine host might
well in experimentally infected pigs.550 Avian-origin maintain a distinct subset of viruses.145-148;154 cited in 146]
H10N7 viruses, isolated from outbreaks among European Serological studies suggest that infections in pigs are
seals in 2014-2015, likewise appeared to be adapting to acquired sporadically from humans and influenza B
replication in seals.564 Antibodies to H1, H2, H3, H4, H6, viruses are not maintained long-term in swine
H7, H8, H10 and H12 viruses have been found in seals, populations.142,149 Limited animal-to-animal trans-
and in some cases, in sea lions and/or walruses (Odobenus mission has been demonstrated in experimentally
rosmarus).65,147,148,561,565 infected guinea pigs, pigs, pheasants and mallard ducks,
but experimentally infected chickens did not transmit the
Influenza A infections have been reported
virus to uninfected chickens. 149,150,156
sporadically in cetaceans, and H1N3, H13N2 and H13N9
viruses have been isolated from whales.7,65 Sequence Influenza C viruses
analysis of the limited isolates available suggests that Influenza C viruses are maintained in people, but
whales do not maintain influenza viruses, but are infected these viruses have been also been isolated from
from other species, most likely birds.566 Antibodies to pigs.7,9,12,14-16,129,576-578 Experimental infections have been
influenza A viruses have been reported in porpoises. 65 established in hamsters, rats, nonhuman primates, dogs,
Influenza A viruses of uncertain origin in pigs and ferrets, although only the dogs and pigs became
other species ill.577,579-581 Serological evidence of infection has been
found in pigs, dogs and horses.7,142-144,582,583
Serological evidence of infection with influenza A
viruses has been reported occasionally from other Influenza D (influenza C-related livestock virus)
mammals not normally thought to be hosts, such as yak, Influenza D viruses have been isolated from cattle,
sheep, goats, reindeer and deer.65,542 Definitive which are thought to be the primary hosts, and from
identification of the virus source can be difficult in pigs.1,2,4-6,355 These viruses might be able to cause illness in
serological studies, but some of these viruses might have both species, but this is still uncertain. Experimental
come from humans.65 Antibodies to influenza A viruses infections have been established in pigs and ferrets.1
have also been reported in reptiles and amphibians
including snakes, crocodiles, alligators, caimans, toads and Zoonotic potential of influenza viruses
frogs, and influenza A viruses have been detected by RT- To date, zoonotic infections have mainly been caused
PCR in caimans, alligators and crocodiles.65 Some of these by swine and avian influenza viruses.
viruses might have been avian, human and equine Clinical cases caused by H1N1, H3N2 or H1N2 swine
influenza viruses.65 influenza viruses are reported sporadically in humans,7,14,97-
99,159,286,287,293,584-598
and serological evidence suggests that
Bat influenza viruses
mild or asymptomatic infections might occur occasionally
The two known bat influenza viruses (H17N10 and in people exposed to pigs.7,14,159,309,337,599-610 Certain
H18N11) may be genetically isolated from other influenza genotypes of swine influenza viruses might be more likely
A viruses, and there is currently no evidence that they to infect humans. Many recent infections in the U.S. were
have been transmitted to any other species.348

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 Influenza
caused by triple reassortant H3N2 viruses that contained the nations now usually exclude these viruses from
‘M’ gene from 2009 pandemic H1N1 virus.96,98,99 commercial, confinement-raised flocks. 49 However,
The two avian influenza viruses reported most often in LPAI viruses may still be present in backyard flocks, live
clinical cases are the Asian lineage H5N1 HPAI viruses and poultry markets and similar sources, 49 Eurasian lineage
recently, H7N9 LPAI viruses in China.14,54,108,258-264,611 H9N2 LPAI viruses and Asian lineage HPAI H5N1
Illnesses caused by H5N1 viruses are, overall, rare; viruses are currently endemic only in parts of Eurasia,
however, these viruses have been found in poultry and have not been detected in the Americas, Australia or
(including small backyard flocks) for over a decade, New Zealand.53,101,184-190,634-639 However, Asian lineage
resulting in high levels of human exposure. The H7N9 virus HPAI H5N8 viruses have reached North America and
in China might be transmitted more readily to people.453,612 reassorted with local strains to produce new variants,
These two viruses could also be identified more often including some that contain North American lineage
because they tend to cause serious, life-threatening illnesses, neuraminidases (e.g., H5N2, H5N1). 220-222 These viruses
which are more likely to trigger laboratory testing than mild occur in wild birds,221,222 but whether they will persist in
flu symptoms.203,237,258-264,613 As of February 2016, Asian the Americas is still uncertain. The zoonotic H7N9 LPAI
lineage H5N8 viruses are not known to have caused any viruses in China have not been reported from other
clinical cases in people, although four cases caused by regions, with the exception of a few imported cases in
H5N6 viruses have been reported in China since 2014.231-234 travelers.262,263,640,641
Illnesses caused by other subtypes, including H6N1, The influenza viruses of pigs and horses tend to occur
Eurasian lineage H9N2 and multiple H7 and H10 avian wherever their maintenance hosts are found, unless there
influenza viruses, have been reported sporadically in are good control programs to exclude them.40,46,93-95,642 As
people.14,53,54,108,159,235,237, described in the Etiology section, different sets of swine
238,241-244,598,611,614-618
Serological surveys in some highly influenza viruses can be maintained on each continent.
exposed populations suggests the possibility of low level Only a few countries, such as New Zealand, Iceland and
exposure to various HA types found in birds, including H4, Australia, are known to be free from all equine influenza
H5, H6, H7, H9, H10, H11 and H12.236,237,239,240,600,605,609,619- viruses.18,19,42,643 At one time, Florida lineage clade 1
629
Experimental infections with some subtypes (e.g., H4N8, equine influenza viruses circulated in North America and
H10N7 and H6N1), have been established in human clade 2 viruses in the Eastern Hemisphere; however, some
volunteers, and some of these viruses caused mild influenza clade 1 viruses have now also become endemic in
symptoms.237 Europe.336,644,645
Very few human infections have been linked to The H3N8 canine influenza virus has been found, at
species other than birds and swine. One person was least sporadically, in most states in the U.S., but its
infected by an H7N7 (avian origin) virus from a seal,561 distribution appears to be patchy.22,25,26,80,646-649 There is no
and a laboratory technician acquired an H1N1 swine evidence that this virus is currently circulating outside the
influenza virus via an infected turkey herd.287,630 U.S. As of 2016, the H3N2 canine influenza virus has
Serological evidence and one experiment in volunteers been reported from South Korea, China, Thailand and
suggest that humans might be susceptible to equine North America (the U.S.).76,81,83,85-88,650 One study reported
viruses,7 but there is no evidence of recent natural antibodies to H3N8 and H3N2 viruses in a few dogs in
infections.13 A survey from Mongolia found that very few Italy,651 but the significance of this finding remains to be
people had antibodies to H3N8 equine influenza viruses, evaluated, as the serological tests were based on reactivity
despite high levels of exposure to horses.631 No infections with equine H3N8 or swine H3N2 viruses, respectively.
with either canine influenza virus have been reported, and There is relatively little information about some
initial studies from the U.S. and Scotland suggest that influenza viruses. Bat-adapted influenza viruses appear to
humans have little or no seroreactivity to influenza D be common in South America.161 Influenza D viruses have
viruses.1,157 been detected in North America, Asia (China) and Europe
(Italy, France), and might be cosmopolitan in their
Geographic Distribution distribution.1,4-6,355
Human seasonal influenza viruses, including 2009
pandemic H1N1 virus, are cosmopolitan.7,52,632,633 Because Transmission
people travel extensively, similar sets of viruses tend to
Transmission of avian influenza viruses
circulate in all populations. However, this is not
among birds
necessarily the case in animals.
Avian influenza viruses can occur in both the feces
Avian influenza (LPAI) viruses occur worldwide in
and respiratory secretions, although the relative amount of
wild birds, but the circulating viruses can differ between
virus may differ, depending on the specific virus, host
regions, and especially between the Eastern and Western
species and other factors.7,15,50,51,371,652,653 While there are
Hemispheres.7,55,197,200 Many countries maintain a HPAI-
some exceptions (including recent Asian lineage H5N1
free status for all poultry. LPAI viruses were once
HPAI viruses121,413,654,655), influenza viruses are
common in poultry, but control programs in developed
predominantly spread by fecal-oral transmission in aquatic

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 Influenza
birds such as waterfowl.55,121,656 On farms, these viruses influenza B viruses (particularly children with diarrhea,
can spread by both fecal–oral and respiratory routes in but also hospitalized adults),689-693 and in severely ill
poultry, due to the close proximity of the birds. Fomites patients infected with pandemic H1N1 virus, zoonotic
can be important, and flies may act as mechanical H7N9 LPAI viruses in China, and Asian lineage H5N1
vectors.51,54,657,658 One study suggested the possibility of HPAI viruses.694-698 In a few cases, the presence of live
wind-borne transmission of HPAI viruses between virus was confirmed by virus isolation. Whether these
farms,659 but this remains to be confirmed. HPAI viruses viruses (or viral RNA) come from swallowed respiratory
have also been found in the yolk and albumen of eggs secretions or other sources is not known.691,693 However,
from some infected chickens, turkeys and quail.660-666 Asian lineage HPAI H5N1 viruses, which can cause
While these eggs are unlikely to hatch, broken eggs could systemic infections, seem to be able to replicate in human
transmit the virus to other chicks in the incubator. It might intestinal tissues.699 H5N1 viruses were also found in the
be possible for birds to shed LPAI viruses in eggs, but the feces of experimentally infected cats and foxes (but not
current evidence suggests this is very rare, if it occurs at some other species, such as pigs),103,114-117,434 while
all.660,667 minimal intestinal shedding of an H3N2 human influenza
How long birds remain contagious differs between virus was reported in experimentally infected raccoons. 468
species, and varies with the severity of the infection. For A ferret model suggests that some influenza viruses
example, chickens and turkeys infected with HPAI viruses might be transmitted to the fetus when there is high
die very quickly, and so do not shed these viruses very viremia10 and viral antigens and nucleic acids were found
long. Most chickens usually excrete LPAI viruses for a in the fetus of a woman who died of an Asian lineage
week, and a minority of the flock for two weeks, but H5N1 infection.700 However, most influenza viruses
individual birds of some species, including waterfowl, can replicate only in the respiratory tract, and transplacental
shed some LPAI or HPAI viruses for a few weeks in the transmission seems much less likely in these cases. A
laboratory.14,370,408,668-670 recent experiment in pigs found no evidence for
transplacental transmission of swine influenza viruses.701
Transmission of influenza viruses in mammals
In mammals, influenza viruses are transmitted in Acquisition of influenza viruses from
droplets and aerosols created by coughing and sneezing, other species
and by contact with nasal discharges, either directly or on Viruses are usually transmitted to other species during
fomites.7,11,12,15,17,31,46,300,671 Close contact and closed close contact with the host or its tissues, although indirect
environments favor transmission.31,672,673 Influenza viruses contact (e.g., via fomites) is also thought to be
enter the body via the respiratory tract, but there is possible.7,14,53,57,59-61,63,64,90,97-99,103,112,263,293,337,548,584,586,595,596,
599-602,605,607,608,612,702-714
increasing evidence that they may also use the eye as a Most of these viruses probably enter
portal of entry.454,479,481,674,675 While aerosol transmission is the body by the respiratory tract. However, a few H5N1
usually thought to occur only during close contact, swine HPAI virus infections in animals, and rare cases in humans,
influenza viruses have been isolated from air samples have been linked to eating raw tissues from infected
inside densely populated pig barns and immediately birds.57,59-61,63,64,103,112,705,712-714 Feeding experiments provide
outside exhaust fans.676,677 One study also detected small evidence that H5N1 viruses can enter the body orally in
amounts of viral RNA up to 2 km downwind of infected cats, pigs, ferrets, mice, hamsters and foxes, and
barns,676 but another group did not find any viral RNA transmission has been confirmed in cats by direct
outside barns, at a distance of 25 m either upwind or inoculation of the virus into the gastrointestinal
downwind.677 The possibility of local airborne spread was tract.59,63,112,114,115,713,715,716 Similarly, uncooked meat from
suggested during a recent equine influenza epidemic pigs or turkeys might have transmitted swine influenza
among naive horses in Australia.678 viruses to mink during two outbreaks,74,534 although it is
Mammals often begin shedding influenza viruses uncertain whether these animals became infected by
before the onset of clinical signs, but the period of virus ingestion or by contamination of the respiratory mucous
excretion is relatively brief in most cases, and typically no membranes.
longer than 7 to 10 days after infection.11,13,15,17,21,26,30,31, Other routes of virus acquisition have been suggested
41,287,329,679,680
Children and adults can occasionally shed in a few cases. Some experiments suggest that turkeys
human influenza viruses for 10 days or more, and viruses might be more susceptible to intrauterine than respiratory
may be detected for weeks in patients who are severely ill or inoculation of pandemic H1N1 virus,717-719 and accidental
immunocompromised.11,12,30,47,681-688 Viral loads are reported transmission during artificial insemination may have been
to vary widely between people infected with human responsible for some outbreaks.490,504 (Turkeys can also be
influenza viruses, and environmental contamination might infected experimentally by intranasal inoculation.531,720)
differ depending on the viral load.688
Host-to-host transmission of novel influenza
Fecal shedding of influenza viruses has been reported viruses
in mammals, although its significance (if any) is still
Animals or people infected with influenza viruses from
uncertain. Viral RNA was found in the feces of a few
other species may or may not transmit the virus to others.
human patients infected with seasonal influenza A or

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 Influenza
Sustained transmission is a rare event, but limited host-to- demonstrated experimentally in both dogs and cats during
host transmission sometimes occurs.7,65,94,95,236,458,528,561,562 close contact.493;731 cited in 732
These events may be difficult to distinguish from exposure Although one H3N8 equine influenza virus adapted to
to a common source of the virus, or transmission on circulate in dogs, other H3N8 equine influenza viruses do
fomites. not seem to spread readily between dogs.90,549
Limited host-to-host transmission of Asian lineage Experimentally infected cats transmitted some equine
H5N1 HPAI viruses has been reported rarely in humans H3N8 viruses to uninfected cats in one study.77
during close, prolonged contact,708-711 among tigers in one
Survival of influenza viruses in the environment
outbreak at a zoo,61 and experimentally between cats.112
There was no transmission of this virus between small Avian influenza viruses
numbers of experimentally infected dogs and cats.116 or The survival of avian influenza viruses in the
between experimentally infected pigs.106 Nevertheless, a environment may be influenced by the initial amount of
recent analysis of H5N1 outbreaks in Indonesia found virus; temperature and exposure to sunlight; the presence
evidence for limited pig-to-pig transmission within (but of organic material; pH and salinity (viruses in water); the
not between) herds.721 Experimentally infected dogs were relative humidity (on solid surfaces or in feces); and in
able to transmit one Asian lineage H5N2 virus to dogs, some studies, by the viral strain.497,668,733-746 Avian
chickens and cats.66,229,230 Another study found that there influenza viruses survive best in the environment at low
was no dog-to-dog transmission of an Asian lineage H5N8 temperatures, and some studies suggest that they are more
virus in the laboratory, and little or no transmission of this persistent in fresh or brackish water than salt
virus was observed in cats despite virus shedding.435 water.497,668,669,733,734,736,738,740,742,746-749 Some viruses from
In rare cases, limited person-to-person transmission has birds may survive for several weeks to several months or
been suspected for some H7 LPAI or HPAI avian influenza more in distilled water or sterilized environmental water,
viruses, including LPAI H7N9 viruses in China.259,263,612,722- especially under cold conditions.733,734,736-738 However, the
727 presence of natural microbial flora can considerably
Transmission typically occurred to family members in
close contact, 259,263,612,722-727 but one H7N9 case was thought reduce this period, resulting in persistence in water for
to have been acquired nosocomially in the hospital.728 There only a few days (or less, in some environments) to a few
was no evidence that close contacts became infected in a weeks.497,737-739,750 Other physical, chemical or biological
number of other avian influenza cases,616,722,729,730 although factors in natural aquatic environments may also influence
seroconversion to some avian viruses may be unreliable in viability.737,738,749,750 In cold climates, freeze-thaw cycles
people.614,724 H9N2 viruses did not seem to spread readily can inactivate influenza viruses, potentially reducing long-
between pigs in one experiment, although these animals did term survival.741
shed virus.252 In feces, some anecdotal field observations stated that
People have occasionally transmitted swine influenza LPAI viruses can survive for at least 44 or 105 days, under
viruses to family members or other close contacts, and a unspecified conditions.733 Under controlled laboratory
limited outbreak occurred on a military base in the 1970s; conditions, LPAI or HPAI virus persistence in feces
however, most cases of swine influenza seem to be ranged from < 1 day to 7 days at temperatures of 15-35°C
acquired by direct contact with pigs.7,14,97- (59-95°F), depending on the moisture content of the feces,
99,286,293,584,586,592,595,702,706,707 protection from sunlight and other
In contrast, swine influenza
viruses transmitted to turkeys can propagate within this factors.497,669,740,743,747,748,751 At 4°C (39°F), some viruses
species.171 survived for at least 30-40 days in two studies,497,747 but
Not surprisingly, the 2009 pandemic H1N1 virus can they remained viable for times ranging from less than 4
cause outbreaks in pigs.93,95,298-300,309,322,326,499-502 One study days to 13 days in two recent reports.740,748 On various
found no evidence for dog-to-dog transmission of this solid surfaces and protected from sunlight, viruses were
virus in the laboratory, although some animals did shed the reported to persist for at least 20 days and up to 32 days at
virus.493 Other studies did observe limited transmission of 15-30°C (59-86°F);669 and for at least 2 weeks at 4°C if the
pandemic H1N1 virus between dogs (although it was relative humidity was low;740 but also for less than 2 days
sporadic and inefficient) or between cats (contact cats on porous surfaces (fabric or egg trays) or less than 6 days
seroconverted but did not become ill).523,530 The length and on nonporous surfaces at room temperature. 752 Virus
size of one pandemic H1N1 outbreak at a cat colony, and survival was longer on feathers than other objects in two
the timing of infections, suggested the possibility of cat-to- reports: at least 6 days at room temperature in one study,752
cat transmission, although the human caretaker was and 15 days at 20°C (68°F) and 160 days at 4°C in another
thought to be the original source of the virus. 521 Animal- report.748. Some viruses survived for up to 13 days in soil
to-animal transmission of pandemic H1N1 virus also (4°C),740 for more than 50 days (20°C) or 6 months (4°C)
appeared possible in a few clustered cases in cats, cheetahs in poultry meat (pH 7),735 and for 15 days in allantoic fluid
and ferrets, but concurrent exposure to an infected human held at 37°C (98.6°F).745 Exposure to direct sunlight
was equally plausible.67,70,71,520,522 Animal-to-animal greatly reduced virus survival.740 Environmental sampling
transmission of H3N2 human influenza viruses was in Cambodia suggests that virus survival in tropical

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 Influenza
environments might be brief: although RNA from Asian populations in the context of disease control.51 The
lineage H5N1 HPAI viruses was found in many samples incubation period for mammalian influenza viruses is
including dust, mud, soil, straw and water, live virus could often 1-3 days, although some cases may take a few days
only be isolated from one water puddle.753 longer to appear.7,13,15-21,25,26,76,82,83,330,530,533,557,679,680,715,763
In particular, incubation periods up to a week have been
Mammalian influenza viruses
reported in some dogs and cats infected experimentally
Human influenza A viruses may also survive for with H3N2 canine influenza viruses.76,82,83,557
months in cold (4°C) water under laboratory conditions,
although infectivity was lost in approximately 2 weeks or Clinical Signs
less at 35°C (95°F).754 Like avian influenza viruses, their
survival was influenced by salinity.754 At room temperature, Highly pathogenic avian influenza
live human influenza viruses could not be recovered from a HPAI viruses usually cause severe illness in chickens
wide variety of surfaces after 24-48 hours, with recovery and turkeys, and few birds in infected flocks survive. 7,51,405
from porous surfaces often lasting less than 8-12 hours.755- Decreased feed and water intake, with other nonspecific
758
Survival on wooden surfaces differed between studies, systemic, respiratory and/ or neurological signs (e.g.,
with one study reporting prolonged survival between 48 depression, edema and cyanosis of unfeathered skin,
hours and 60 hours.758 One group reported that human diarrhea, ecchymoses on the shanks and feet, coughing)
influenza viruses remained viable for up to 3 days on Swiss are common clinical signs, but no signs are
banknotes, and for as long as 8-17 days if the viruses were pathognomonic, and sudden death can also be seen.15,49-
51,54,175,405,406,411,664,764-767
in nasopharyngeal secretions.759 Because a virus can be defined as
In one study, swine influenza viruses were inactivated highly pathogenic based on its genetic composition alone,
in untreated pig slurry in 1-2.5 hours at 50-55°C (122- HPAI viruses may rarely be isolated from chicken or
131°F), two weeks at 20°C (68°F), and 9 weeks at 5°C turkey flocks that have mild signs consistent with low
(41°F).760 pathogenic avian influenza.50,182
Infections with HPAI viruses may be asymptomatic,
Disinfection mild or severe in other domesticated and wild (or captive
Influenza A viruses are susceptible to a wide variety wild) avian species, including gallinaceous birds other
of disinfectants including sodium hypochlorite, 60% to than chickens and turkeys. 7,15,48,50,51,55,62,101,120,121,365,
388,390,392,393,395-397,405-408,422,768-770
95% ethanol, quaternary ammonium compounds, Domesticated waterfowl
aldehydes (glutaraldehyde, formaldehyde), phenols, acids, tend to have minimal or mild signs, but respiratory signs
povidone-iodine and other agents.11,15,21,158,733,761 Common (e.g., sinusitis), diarrhea, corneal opacity, occasional
household agents, including 1% bleach, 10% malt vinegar neurological signs, and somewhat increased mortality
or 0.01-0.1% dishwashing liquid (washing up liquid), as may be seen, and some Asian lineage H5N1 HPAI
well as antimicrobial wipes, were found to destroy the viruses can cause severe acute disease with neurological
viability of human influenza viruses, although hot water signs and a high mortality rate. 15,51,52,109,119-
121,395,413,414,771,772
(55°C; 131°F) alone was ineffective for rapidly These H5N1 viruses have caused
eliminating viruses.762 Influenza A viruses can be sudden deaths and severe systemic, respiratory and/or
inactivated by heat of 56-60°C (133-140°F) for a neurological signs in some free-living and captive wild
minimum of 60 minutes (or higher temperatures for birds, although mild signs or subclinical infections are
shorter periods), as well as by ionizing radiation or also possible.62,109,120,121,365,392,393,395-398,414,425,451,773-775
extremes of pH (pH 1-3 or pH 10-14).15,158,669,733,761 The H5N8 HPAI viruses and their reassortants may likewise
disinfectant and heat susceptibility of influenza B and C be associated with a wide range of outcomes in wild
viruses has not been examined extensively, but it is birds: these viruses have been found in sick, dead and
probably similar.12 apparently healthy waterfowl, and sick or dead birds in
several other orders including raptors. 219,221,224,226,386,387,
401,402,404
Infections in Animals
Low pathogenic avian influenza
[Note: for more detailed information on avian, swine,
equine and canine influenza, please see individual LPAI viruses (including the Chinese H7N9 viruses)
factsheets on these diseases at http://www.cfsph. usually cause subclinical infections or mild illnesses in
iastate.edu/DiseaseInfo/factsheets.php.] poultry and other birds.48,51,264,424,450,451 In chickens and
turkeys, there may be decreased egg production and egg
Incubation Period quality, respiratory signs, lethargy, decreased feed and
The incubation period for influenza is short in all water consumption, or somewhat increased flock mortality
species. In poultry, it can be a few hours to a few days in rates.49,51,163,664,776-784 Illnesses exacerbated by factors such
individual birds, and up to 2 weeks in the flock.15,49,51 A as concurrent infections or young age can be more
21-day incubation period, which takes into account the severe.48-50,785 Wild birds have few or no obvious clinical
transmission dynamics of the virus, is used for bird signs,55,786 although subtle effects (e.g., decreased weight
gain, behavioral effects or transient increases in body

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 Influenza
temperature) have been described in some free-living (ferrets and mice). However, H9N2 LPAI viruses have
birds.188,787,788 been found occasionally in sick pigs in China, and pigs
The H9N2 viruses currently circulating among poultry inoculated with some H9N2 viruses (but not others) had
in the Eastern Hemisphere can cause significant lesions of pulmonary congestion. 252 Dogs and cats
respiratory signs and malaise in chickens, including inoculated with H9N2 viruses remained asymptomatic in
experimentally infected chickens that are not co-infected two studies,446,447 but dogs in another study had nonfatal
with other pathogens.789,790 Clinical signs have been respiratory signs. 445 Few or no clinical signs were seen in
reported in quail, which are usually mildly affected by cats inoculated with an H7N7 HPAI virus isolated from a
other viruses, and some experimentally infected quail fatal human case, cats inoculated with several LPAI
became severely ill.49,637,791 Some wild birds also viruses from waterfowl, or raccoons experimentally
developed clinical signs after experimental inoculation infected with an H4N8 virus. 461,462,468 The Chinese
with H9N2 viruses.789 (zoonotic) H7N9 LPAI virus caused fever alone in
experimentally infected cynomolgus macaques and no
Avian influenza viruses in mammals clinical signs in miniature pigs. 453 Avian influenza
Asian lineage H5N1 HPAI viruses have caused fatal viruses have occasionally caused outbreaks in marine
disease, as well as milder illnesses or asymptomatic mammals and mink (see separate sections on these
infections, in mammals. A few clinical cases have been species, below).
described, at most, in each species. One group of infected
cats had no clinical signs, but a few other infected cats Swine influenza
were found dead, and one cat developed fever, dyspnea Swine influenza is an acute upper respiratory disease
and neurological signs before it died.57,58,103,712 with coughing and other respiratory signs, and nonspecific
Conjunctivitis and severe or fatal respiratory signs were signs such as lethargy and weight loss.7,14-17,41,642 Some
described in experimentally infected cats. 112,114,116,715,792 outbreaks are more severe than others, and swine influenza
Some captive tigers and leopards exhibited high fever, viruses can circulate in pigs with few or no clinical
respiratory distress and neurological signs before signs.7,14,46,92 Depending on the production system, illness
death,59-61,65,211 while a non-fatal outbreak among large may be seen in certain age groups, while others remain
felids was characterized by lethargy and inappetence asymptomatic.41,793 Concurrent infections with other
without respiratory signs. 62 Fever, respiratory and/or pathogens can exacerbate the clinical signs.14,16,41,793
neurological signs were also reported in a handful of Swine influenza viruses in turkeys and ferrets
cases in other species, including a dog, captive raccoon
Turkeys infected with swine influenza viruses may
dogs, captive palm civets and a wild stone marten. 63-65,122
develop respiratory disease, have decreased egg
Experimental infections in various mammals ranged from
production, or lay abnormal eggs.15,171
subclinical to severe, with systemic and/or respiratory
signs reported in animals that became ill. 106,110,115- During an outbreak caused by a triple reassortant
118,429,434
Experimental infections in dogs tended to be H1N1 swine influenza virus, ferrets developed respiratory
mild or asymptomatic unless the dogs were inoculated by signs, including dyspnea, and some severely affected
a route (e.g., intratracheal) that bypasses natural animals died.173
respiratory defenses. 116,117,429,792 A recent comparative Equine influenza
study also suggested that experimentally infected dogs Equine influenza is an acute respiratory disease,
were much less likely to develop clinical signs than which usually begins with a high fever, followed by a
cats.792 Experimental infections, as well as reports of deep, dry, often paroxysmal cough, serous to
infected herds, suggest that H5N1 HPAI virus-infected mucopurulent nasal discharge, and other respiratory and
pigs usually remain asymptomatic or have only mild nonspecific signs.7,13,15,18,20,21,329,543 Animals with partial
signs.106,115,430,431,721 One H5N1 virus was isolated from immunity can have milder, atypical infections,20 while
donkeys during a respiratory disease outbreak in Egypt, young foals without maternal antibodies may develop
although a causative role is uncertain, and antibodies to severe viral pneumonia.7,20,21 Some cases may be
these viruses were detected in healthy donkeys and complicated by secondary bacterial infections, sequelae
horses.122,431 such as chronic bronchitis, or rare complications such as
Relatively little is known, at present, about other neurological signs, myocarditis, myositis or limb
Asian lineage H5 viruses. One dog naturally infected with edema.13,18,20,21,329,336 Healthy adult horses usually recover
an H5N2 HPAI virus developed nonfatal respiratory signs, within 1-3 weeks, although the cough may persist
and similar signs occurred in dogs and one cat inoculated longer.7,13,18,21,329 Convalescence can take months in
with this virus.66,229,230,445 No clinical signs were seen in severely affected animals.18
dogs experimentally infected with an Asian lineage H5N8
HPAI virus, while cats had mild and transient signs, Equine influenza viruses in other animals
including fever and marginal weight loss.435 Mild to severe respiratory signs have been reported in
There are few reports of animals infected with other dogs naturally infected with H3N8 equine influenza
avian viruses, except as animal models for human disease viruses.89,90,544,548 while experimental inoculation resulted

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 Influenza
in mild or no signs.548,549 One recent H3N8 isolate caused viruses circulating before 2009 usually recovered within 2
respiratory disease in experimentally infected cats, but an weeks, although neonates sometimes became severely ill
older isolate did not make cats ill.77 or died.10,485,487 Similar signs were reported in ferrets
Depression and respiratory signs, including coughing, infected with pandemic H1N1 virus; however, some cases
were reported in pigs infected with an H3N8 equine in adults were severe or fatal.510,511,513,520,522,527
influenza virus in China.547 Streptococcus suis was also Experimental infections differed in severity, with some
recovered from these herds, and hemorrhagic pleuritis was ferrets developing milder respiratory signs or systemic
noted in some animals at necropsy. Another equine H3N8 signs (e.g., lethargy and weight loss, with little
virus did not replicate well in experimentally infected sneezing),525,526 but other studies suggesting that pandemic
pigs.550 One H3N8 virus was recovered from a healthy H1N1 virus might be more virulent in ferrets than other
camel.546 human H1N1 viruses.529
Mild as well as severe or fatal illnesses have been
Canine influenza (H3N8)
seen in cats infected with pandemic H1N1 virus.67-
H3N8 viruses tend to cause a relatively mild illness, 71,514,517,521
The clinical signs included anorexia, lethargy,
with a low fever alone, or fever followed by malaise, a upper or lower respiratory signs ranging from sneezing
persistent cough and other respiratory signs.22-27,79 The and nasal discharge to dyspnea, and concurrent issues such
clinical signs may last for up to 3 weeks regardless of as dehydration. Fever was not reported in some cases at
treatment Secondary bacterial infections seem to be presentation. Some cats remained ill for several weeks.
common, resulting in mucopurulent nasal discharge and One cat that died had evidence of myocarditis in addition
other signs.25 Pneumonia or bronchopneumonia can to lung involvement at necropsy, but whether this was a
develop in more severe cases, but this has generally been pre-existing condition or a consequence of the viral
associated with concurrent bacterial or mycoplasmal infection is not known.68 Experimentally infected cats
infection.22-24,26,28,79 Peracute deaths with evidence of became mildly to moderately ill with lethargy, loss of
hemorrhages in the respiratory tract occurred during the appetite and respiratory signs.530 Two clinical cases
initial outbreaks among racing greyhounds; however, this reported in dogs were characterized by fever and
syndrome does not seem to be prominent in pets.24,27 radiological evidence of pneumonia, while a third dog had
Canine influenza (H3N2) only a severe cough, with mild depression and
Like other influenza viruses, H3N2 canine influenza anorexia.518,523 Mild fever, occasional mild coughing, and
viruses cause respiratory signs.76,81-84,86,87,554,556 Most nasal discharge were the only signs in experimentally
reports from Asia have described severe or relatively infected dogs.523
severe outbreaks and case series in dogs, or dogs and Pandemic H1N1 virus also caused respiratory signs in
cats.76,81-84,86,87,554,556 Deaths were common in several of captive wild species including cheetahs, giant pandas, a
these reports, but coinfections with other pathogens may black-footed ferret, an American badger and a
have sometimes played a role. Experimentally infected binturong.67,72,73 Some cases, including those in four
cats developed elevated temperatures, lethargy and cheetahs, the badger and binturong were severe, although
respiratory signs,556 while experimentally infected dogs the cheetahs recovered with supportive care including
had fever, respiratory signs and severe pathology.81,82,84,794 antibiotics.67,72 The pandas (which were the only animals
Serological surveys of healthy dogs and cats in Asia have to receive antiviral drugs) and black-footed ferret also
suggested that some animals might have mild or recovered.72,73 Pandemic H1N1 virus was detected in wild
asymptomatic infections and recover without striped skunks found dead with severe mixed bacterial
treatment.86,88,246,443,555,558,795-799 There are currently no bronchopneumonia, thought to be secondary to viral
published articles describing the outbreaks in the U.S., but infection, and concurrent Aleutian disease virus
informal reports suggest that most cases have been infection.519 These skunks came from a mink farm where
characterized by relatively mild upper respiratory signs, many of the animals had nasal discharge; however, the
with few deaths.800 clinical signs in the mink were not investigated. Another
Ferrets do not seem to be very susceptible to the outbreak of respiratory disease in mink (see below) was,
canine H3N2 virus, although some experimentally infected however, confirmed to be caused by pandemic H1N1. 801
animals developed mild signs including sneezing.556,559 In pigs, infections with pandemic H1N1 virus are
Experimentally infected guinea pigs remained usually mild and resemble swine influenza.275,322,502,802-812
asymptomatic, but had lung lesions.560 Decreased egg production and quality were the only
significant signs in most turkeys,490,504-509 although one
Pandemic H1N1 and other human influenza A flock coinfected with Pasteurella multocida (fowl cholera)
virus infections in animals developed mild clinical signs and had slightly increased
Ferrets that are naturally infected with seasonal mortality.509
human influenza viruses may develop a febrile respiratory Horses experimentally infected with one human
disease with anorexia, depression, sneezing, nasal influenza virus (H3N2 ‘Hong Kong’) developed a mild
discharge and coughing.10,485,486 Adult ferrets infected with

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 Influenza
febrile illness.7 Raccoons inoculated with human H3N2 mild respiratory signs of sneezing and serous nasal
viruses remained asymptomatic.468 discharge; however, the clinical signs were overall mild or
absent.156 In a recent study, some experimentally infected
Influenza viruses in mink
pigs similarly developed a transient fever, lasting 1-3 days,
Outbreaks in mink caused by H10N4 and H9N2 and minor lesions consistent with influenza were detected
avian influenza viruses, H3N2 and H1N2 swine influenza at necropsy.149 Ponies inoculated with influenza B viruses
viruses and pandemic H1N1 virus were characterized by sometimes had a fever for 1-3 days, and one animal
respiratory signs of varying severity. 7,65,74,75,441,534,801 developed nonspecific, self-limiting signs of illness
There were few or no deaths during some of these including sweating, listlessness and unusually heavy
outbreaks, but pneumonia and an increased mortality rate breathing.571 Influenza B infections have also been
were reported in others, particularly during the H10N4 reported in some stranded seals.65
avian influenza outbreak, and in kits and on farms where
the mink were co-infected with other pathogens. Mink Influenza C
infected with an H1N2 swine influenza virus were co- Respiratory signs were reported in pigs and dogs
infected with hemolytic E. coli, and developed severe inoculated with influenza C viruses. The pigs had nasal
respiratory disease with hemorrhagic bronchointerstitial discharge and slight dyspnea, without fever.577 Two pigs
pneumonia.534 The hemorrhagic pneumonia and high recovered quickly, but two others had clinical signs for 10
mortality were attributed to the secondary bacterial days. The dogs developed conjunctivitis and nasal
component.534 Mink that were experimentally infected discharge, which persisted in most animals for more than
with H1N1 or H3N2 human influenza viruses, H1N1 10 days.579 Experimentally infected rats, hamsters, ferrets
swine influenza virus, H3N8 equine influenza virus, and and nonhuman primates did not become ill.580,581
H3N8 and H4N6 avian influenza viruses remained
Influenza D (Novel livestock-associated influenza
asymptomatic despite shedding virus. 65 Mink inoculated
C virus)
with an H9N2 avian influenza virus developed mild
signs.441 Whether influenza D viruses cause any illnesses is
still uncertain. These viruses have been recovered from
Influenza in marine mammals apparently healthy cattle; however they were also found in
Influenza A (avian origin) viruses have been some clinical samples from cattle with respiratory signs. 2,4-
6,355
associated with outbreaks of pneumonia or mass mortality Influenza D viruses were originally detected in a herd
in seals.7,172,563,564,566,763 The clinical signs in some of pigs exhibiting respiratory signs that resembled
outbreaks (including outbreaks in well-fed captive influenza, and they have since been found in other pigs
animals) included weakness, incoordination, dyspnea, and with similar signs.1,5 Neither pigs nor ferrets developed
in some cases, subcutaneous emphysema of the clinical signs or gross lesions after experimental
neck.65,561,763 Some animals had a white or bloody nasal inoculation.1
discharge. Experimental infections with some viruses were
milder or asymptomatic, suggesting that co-infections may Post Mortem Lesions Click to view images
have increased the severity of the illness.65 Clinical signs Highly pathogenic avian influenza in birds
and mortality were reported only in harbor seals (Phoca The lesions in chickens and turkeys are highly
vitulina) during one H10N7 outbreak, although gray seals variable and resemble those found in other systemic avian
(Halichoerus grypus) also had serological evidence of diseases.52,815 Classically, they include edema and cyanosis
infection.564,813 of the head, wattle and comb; excess fluid (which may be
An influenza virus was isolated from a diseased pilot blood-stained) in the nares and oral cavity; edema and
whale, which had nonspecific signs including extreme diffuse subcutaneous hemorrhages on the feet and shanks;
emaciation, difficulty maneuvering and sloughing skin. 763 and petechiae on the viscera and sometimes in the
Whether this virus was the cause of the disease or an muscles.49,51,815 There may also be other abnormalities,
incidental finding is uncertain.528 Other influenza viruses including hemorrhages and/or congestion in various
were isolated from whales that had been hunted, and were internal organs, as well as severe airsacculitis and
not linked with illness.814 peritonitis (caused by yolk from ruptured ova). 51 However,
Influenza B the gross lesions in some outbreaks may not fit the
classical pattern,815 and birds that die peracutely may have
There is little information about the effects of
few or no lesions.49,51,815 Lesions reported from fatal cases
influenza B viruses, if any, on mammals other than
in other species of birds vary.393,395,396,425,816
humans. One early field report suggested that an
influenza-like outbreak in swine might have been caused Low pathogenic avian influenza and pandemic
by viruses acquired from humans, based on serology and H1N1 viruses in birds
the timing of the illness, which occurred concurrently with Poultry infected with LPAI viruses may exhibit
a human influenza B outbreak.155 cited in 156 A follow-up rhinitis, sinusitis, congestion and inflammation in the
study found that pigs inoculated with influenza B virus trachea, but lower respiratory tract lesions such as
developed elevated temperatures, and some animals had

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 Influenza
pneumonia usually occur only in birds with secondary Diagnostic Tests
bacterial infections.49,51 Lesions (e.g., hemorrhagic
ovaries, involuted and degenerated ova) may be observed Influenza A viruses
in the reproductive tract of laying hens, and the presence Avian influenza viruses, their antigens and nucleic
of yolk in the abdominal cavity can cause air sacculitis and acids can be detected in respiratory and/or intestinal
peritonitis.51 A small number of birds may have signs of samples (e.g., cloacal swabs) from birds.50 Samples from
acute renal failure and visceral urate deposition.49 various internal organs are also tested in dead birds
Reproductive lesions, with peritonitis in some cases, suspected of having HPAI.50,51 Respiratory samples are
were the only lesions reported in turkeys infected with usually taken from mammals (e.g., nasal or
pandemic H1N1 virus.504 nasopharyngeal swabs from living animals, or lung tissue
samples collected at necropsy).15-17,26,543,818
Influenza lesions in mammals Virus isolation is useful for the characterization of
The major lesions caused by the influenza viruses of influenza viruses, and can be used in diagnosis, although
mammals are usually lung consolidation and/or faster and simpler techniques such as RT-PCR tend to be
pneumonia, or upper respiratory tract involvement alone employed in most clinical cases. Avian influenza viruses
in milder cases.16,19-21,24,26-28,41,65,172,561,679,680,763,817 are isolated in embryonated eggs,50 while mammalian
Concurrent bacterial infections, common in naturally influenza viruses can be isolated in embryonated chicken
infected animals, can result in more extensive damage to eggs or cultured cell lines.14,17,543,818 Both eggs and cell
the lungs.14,41 Lower respiratory tract lesions were cultures can be used to maximize the recovery of some
reported in some animals infected with 2009 pandemic mammalian viruses.26,543 Virus shedding is usually brief in
H1N1 virus.68,70,533 mammals, and respiratory samples should be collected
Some viruses have caused severe illnesses resulting very soon after the onset of clinical signs.17,24,329,543,679,680
in hemorrhagic lesions in the lungs. Hemorrhagic Isolation of the H3N8 canine influenza virus from live
pneumonia occurred in fatal cases caused by the H3N8 dogs can be difficult.24,679,680 A virus detected in culture
canine influenza virus in greyhounds, although this can be identified as influenza A with agar gel
syndrome seems to be uncommon in other dogs infected immunodiffusion (AGID), antigen-detection ELISAs or
with this virus. 26-28,79,817 Severe hemorrhagic other immunoassays, or by a molecular test such as RT-
bronchointerstitial pneumonia was reported in most fatal PCR.49,50 Viruses can be subtyped with specific antisera in
cases of canine H3N2 influenza in dogs in Asia (although hemagglutination and neuraminidase inhibition tests, by
few necropsies were done), 794 and dogs inoculated with RT-PCR, or by sequence analysis of the viral HA and NA
this virus had pneumonia with consolidation, edema and genes.14,50,818 Genetic tests to identify characteristic
hemorrhages.81,82,84,794 Hemorrhagic lesions were also patterns in the HA (at its cleavage site) and/or virulence
found in the respiratory tract and intestinal serosa of two tests in young chickens are used to distinguish LPAI
cats that died during a pandemic H1N1 outbreak in a cat viruses from HPAI viruses.50,51
colony,521 although more typical influenza lesions were RT-PCR assays are often used to detect influenza
reported from other cases in cats. 68,70 Influenza-related viruses in clinical samples.13,14,17,18,50,51,329,543,818-820 Real-
pneumonia in harbor seals is characterized by necrotizing time RT-PCR is the method of choice for the diagnosis of
bronchitis and bronchiolitis and hemorrhagic avian influenza in many laboratories,50,51 and it is also one
alveolitis.763 The lungs were hemorrhagic in a whale of the two most reliable techniques for diagnosing H3N8
infected with influenza virus, 763 although the lesions canine influenza (the other is serology).26,821,822 Viral
could not be definitively attributed to this virus. 528 antigens can be identified in clinical samples with various
Avian H5N1 influenza viruses in mammals tests (e.g., ELISAs in various species; immune-
histochemistry or immunofluorescent techniques; and
Asian lineage H5N1 HPAI viruses can cause systemic
other individual tests validated for a species).13,14,17,18,26,
lesions as well as pulmonary lesions in some animals. 50,329,543,818,819
The sensitivity and uses of these tests can
Gross lesions reported in some cats and other felids
differ between species.26,50,822
included pulmonary congestion and/or edema, pneumonia,
hemorrhagic lesions in various internal organs, and in Serological tests may be used for diagnosis and/or
some cases, other lesions such as multifocal hepatic other purposes. In birds, serology can be valuable in
necrosis, or cerebral, renal and splenic surveillance; however, it is not very useful for diagnosing
congestion.57,58,60,103,112,114,715 Bloody nasal discharge, HPAI infections in highly susceptible birds, which usually
severe pulmonary congestion and edema, and congestion die before any antibodies develop. Serological tests used
of the spleen, kidney and liver were reported in a naturally in poultry include AGID, hemagglutination inhibition (HI)
infected dog.63 Pulmonary lesions including interstitial and ELISAs.50 AGID tests and ELISAs that detect
pneumonia have been noted in some experimentally conserved influenza virus proteins can recognize all avian
infected pigs,106 while others had mild to minimal gross influenza subtypes, but HI tests are subtype specific. In
lesions.115 mammals, influenza is sometimes diagnosed
retrospectively with a rising antibody titer in paired serum

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 Influenza
samples.16,18,24,329,543,821,823 Single tests are occasionally requirements for the various influenza viruses differ
helpful when pre-existing titers are absent or uncommon between countries, but HPAI viruses are generally
(e.g., canine influenza in some dog populations). 26,823 reportable. Veterinarians who encounter or suspect a
Serological tests employed in mammals include HI, and in reportable disease should follow their national and/or local
some species, other tests such as single-radial hemolysis, guidelines for informing the proper authorities (state or
ELISAs and virus neutralization.13,14,16,17,24, federal veterinary authorities in the U.S.). Unusual
26,543,818,821
Cross-reactivity between influenza viruses can mortality among wildlife should also be reported (to state,
sometimes be an issue. In addition, some studies have tribal or federal natural resource agencies in the U.S.826)
found that mammals infected by viruses adapted to other
Prevention
species may fail to develop antibodies to the viral HA,
despite having antibodies to other viral proteins, such as Vaccines
the viral nucleoprotein.465,550 Vaccines are available for avian, swine and equine
Tests that can distinguish infected from vaccinated influenza viruses, and in some countries, for H3N2 or
animals (DIVA tests) may be used in some surveillance H3N8 canine influenza viruses.7,13,15-18,26,49,51,329,336,543,827,828
programs in birds.50,165,824 DIVA tests are rarely available A poor match between the vaccine and virus can
for mammalian influenza viruses; however, one ELISA compromise protection.297 In pigs, some combinations of
was used with a canarypox-vectored vaccine during the swine influenza vaccines and poorly matched challenge
2007-2008 equine influenza virus eradication campaign in viruses were reported to exacerbate disease, at least in a
Australia.330,336,678 Another DIVA ELISA, based on the laboratory setting.91,829-831
NS1 influenza protein, has been suggested for possible use In birds, vaccine use may be complicated by the need
with inactivated vaccines in horses.329 to keep commercial flocks free of LPAI viruses, and to
Influenza D viruses quickly recognize the introduction of HPAI viruses into a
country. Although routine vaccination can suppress
Diagnostic testing for influenza D viruses is not yet clinical signs, birds may still become infected and shed
standardized. Some investigators have reported that these viruses. This can prevent infected flocks from being
viruses can be readily isolated in two mammalian cell recognized if good surveillance programs are not used
lines.1,355 Other researchers had variable success in simultaneously.51,832-834 In addition, vaccination programs
recovering influenza D viruses with mammalian cell lines.4,5 can place selection pressures on influenza viruses, which
RT-PCR has been employed by several groups and serology may encourage the evolution of vaccine-resistant
was used during the initial studies in the U.S.1,2,4-6 isolates.835-838 While avian influenza vaccines are used
Treatment routinely in some regions, other countries (including the
U.S.) restrict their use.49,50 In some cases, avian influenza
Mammals with influenza are usually treated with
vaccines may be used as an adjunct control measure
supportive care and rest.13,16,18,49 Antibiotics may be used
during an outbreak (in conjunction with surveillance and
to control secondary bacterial infections.13,16,18,23,49
movement controls), or to protect valuable species such as
Antiviral drugs used in humans are not generally given to
zoo birds.50,561,832
animals, although ferrets infected with human influenza
viruses have been treated with amantadine.485 (The Influenza vaccines for animals are changed
usefulness of this drug will vary with the antiviral periodically to reflect the current subtypes and strains in
resistance patterns of the circulating strains, see Human the area, although antigenic drift tends to be lower than in
Treatment section below.) Antiviral drugs (oseltamivir) human influenza viruses.13-15,329,336. A multi-country
were used in captive giant pandas infected with pandemic surveillance program for equine influenza viruses
H1N1 virus,73 and some authors have speculated that they recommends changes in vaccine strains.330,828 Although
might be of use in valuable horses.52,825 One issue with such programs do not currently exist for swine influenza
antiviral drugs is that the brief period when viruses are viruses, surveillance has increased since the 2009-2010
most susceptible (48 hours) has often passed by the time human pandemic. Swine influenza viruses in North
the animal is seen.26 The potential for influenza viruses to America have recently become very diverse,40,297 making
develop resistance to these drugs is an additional concern. vaccination in this area a challenge.
Poultry flocks infected with HPAI viruses are Other preventive measures
depopulated (this is generally mandatory in HPAI-free Biosecurity measures help prevent influenza viruses
countries) and not treated. from being introduced into a flock, herd or exhibit. In
addition to routine hygiene and sanitation, some sources of
Control infection to consider are contact with susceptible wild
Disease reporting species (e.g., wild birds for avian influenza, and wild or
A quick response is vital for containing outbreaks in feral swine for swine influenza), fomites, drinking water,
regions that are free of a virus, and in some cases, for raw feed (e.g., pork or poultry fed to mink), and humans
minimizing the risk of zoonotic transmission. Reporting who may be infected with viruses transmissible to
animals.15,39,52,54,57,59-61,63,67-69,74,103,112,300,510-520,534,561,733,839-841

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 Influenza
Management measures such as all-in/all-out production HPAI viruses cause severe illness even in waterfowl, and
can help prevent the introduction of viruses in new the introduction of these viruses may be heralded by
animals.14,15,46,52,54,287 Isolation of newly acquired animals unusual deaths among wild birds (e.g., swans in Europe
(or animals returning to a facility) and testing before and recently crows in Pakistan).7,14,52,62,120,121,392,393,398,845,846
release also decreases the risk that the rest of the herd or Some H5N1 HPAI outbreaks, such as one at Qinghai
flock will become infected.18,300,329 Isolating infected Lake, China in 2005, have killed thousands of wild
animals may help reduce transmission within a facility birds.847. Wild bird deaths have also been associated with
during an outbreak,13,840 and quarantines (voluntary self- some Asian lineage H5 reassortants, such as H5N8
quarantine or government-imposed) reduce transmission viruses, in Asia and North America, although these viruses
between premises.13,15 Management measures, such as have also been detected in apparently healthy wild
resting horses, can help decrease the severity of the birds.219,221,224,
illness.7,13,14,46,287 226,386,387,401,402,404

Preventive measures for pets include awareness of Mammals

potential susceptibilities (e.g., human seasonal influenza
Mammalian influenza viruses differ in prevalence in
viruses in ferrets, pandemic H1N1 in cats, Asian lineage
their host species. Some viruses are very common. For
H5N1 in multiple species) and, to the extent practical,
example, a number of studies have reported that
avoidance of close contact with the source of the infection.
approximately 20-60% of domesticated pigs have
Eradication antibodies to swine influenza viruses, with lower rates in
HPAI viruses are normally eradicated by depopulating feral swine and wild boar.7,14,46,93,142,300,503,538,818 In contrast,
infected flocks, combined with other measures such as North American H3N8 canine influenza viruses are
movement controls, quarantines and perhaps vaccination. currently uncommon in pets, possibly because virus
Infected swine herds can be cleared of influenza viruses by shedding is low and transmission between these animals is
depopulation46,287 or management measures.300 Elimination inefficient.26,848,849 However, these viruses are more
of a mammalian influenza virus from an entire country is prevalent where dogs are in close contact, such as kennels
unusual; however, Australia successfully eradicated an and animal shelters.849-851
introduced equine influenza virus with quarantines, During outbreaks, influenza viruses can spread rapidly
movement controls, vaccination, and both serological and in fully susceptible, exposed populations. Morbidity rates
virological testing (including the use of an ELISA that of 60-90% or higher have been reported in some naïve
could distinguish vaccinated from infected horses).678,842 horse populations during equine influenza
epidemics,7,13,20,21 and infection rates may approach 100%
Morbidity and Mortality during canine H3N8 virus outbreaks in kennels.25,26
Birds In healthy mammals, uncomplicated infections with
Influenza virus exposures and shedding patterns among host-adapted equine and swine viruses are usually
wild birds are complex and likely to reflect their exposure to associated with low mortality rates and rapid recovery
different habitats, as well as gregariousness and other social from the acute stage of the illness, although signs such as a
factors, and pre-existing immunity.56,361 The reported cough may linger.7,13-21,329 However, the severity of the
prevalence of LPAI viruses among wild birds ranges from illness can vary with the dose and strain of virus, and host
<1% to more than 40%, typically with much higher rates in factors such as young age, pre-existing immunity (or
birds from aquatic environments than terrestrial maternal antibodies), stressors such as transport and
species.188,193,356,358-360,363,366,367,372,373,377,843,844 Currently, concurrent illnesses, and secondary bacterial
surveillance suggests that carriage of H5N1 HPAI viruses in infections.14,16,18,20,39-42,543 More severe clinical signs have
wild bird populations without unusual mortality events is also been reported in pregnant mares close to parturition. 42
rare.428,845 The prevalence of influenza viruses in poultry The H3N8 canine influenza virus has followed this
differs between nations, but confinement-raised commercial pattern of generally high morbidity and low mortality,
poultry in developed countries are generally free of LPAI except during the initial outbreaks in racing greyhounds,
and HPAI viruses.49 when severe and fatal cases were common.22-27 Several
LPAI viruses usually cause mild illnesses or papers have reported severe H3N2 canine influenza virus
asymptomatic infections in poultry, including chickens, outbreaks in Asia, with a case fatality rate of 50% in two
turkeys and ducks, but the outbreak can be more severe small case series in pets; case fatality of 25% in dogs and
when there are concurrent infections or other exacerbating 40% in cats during one explosive outbreak at an animal
factors.48,50,51 Chicken and turkey flocks infected with shelter; and morbidity and mortality rates of 77% and
HPAI viruses have high cumulative morbidity and 23%, respectively, in dogs, and 47% and 22%,
mortality rates, which may approach 90-100%.51,53 HPAI respectively, in cats, at another animal shelter.76,81,86,554
viruses can cause mild or severe disease in other species, However, clinical cases were reported to be relatively mild
and domesticated or wild waterfowl are often mildly after an H3N2 virus was introduced to the U.S., and the
affected.370,405-412 However, some Asian lineage H5N1 case fatality rate was low: news reports indicated that, as
of May 2015, there were approximately 8 confirmed

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 Influenza
deaths in more than 1500 cases in dogs.800 The reason for that dogs and cats may be infected with other human
this discrepancy is not clear, although serological studies influenza viruses. A number of surveys have found that only
of dogs and cats in Asia also suggest that a significant a few animals (< 5%) had antibodies to various human
number of animals there might have been infected without seasonal influenza viruses including pandemic H1N1 virus,
severe clinical signs.85,86,88,246,443,555,558,795-799 and a few reported no reactivity, but others reported
seroreactivity of up to 44% for some viruses (and rarely,
Viruses acquired from other species
even higher), depending on the animal population, virus and
Few generalizations can be made about influenza test used.333,492,732,796,798,853-863;731 cited in 732
viruses acquired from other species; however, pigs seem to
Only a few instances of cross-species transmission have
be infected fairly regularly by viruses from birds and
been reported in horses, but an avian H3N8 virus resulted in
humans, often with only minor consequences even when the
a 20-35% mortality rate when it was introduced into horses
virus belongs to the Asian lineage of H5N1 HPAI
in China, with little or no mortality in subsequent years.13,19
viruses.7,14,15,27,46,94,95,104,106,109,115,129,142,144,252,430,431,438,452,524,533,
576,721,852
The consequences of infection with these H5N1
viruses have varied widely in other mammals. Infections in Infections in Humans
cats and other felids ranged from asymptomatic to fatal,
[Note: for more detailed information on zoonotic
while dogs generally appear to be less severely affected than
influenza caused by avian and swine influenza viruses,
cats, and sporadic deaths have been reported in other species
please see individual factsheets on these animal diseases]
such as raccoon dogs, palm civets, and a mink,57-
65,103,110,112,114-117,211,246,429,712,792
Experimental infections in Incubation Period
various species, likewise, varied from subclinical or mild to The incubation period for seasonal human influenza,
severe and fatal,110,115,116,429,434 and some studies have including infections caused by pandemic H1N1 virus, is
reported antibodies to H5 viruses in some cats, dogs, horses, short, with most cases appearing in 1-4
donkeys and pigs tested in Asia or days.7,11,31,37,47,864,865 Most zoonotic infections caused by
Egypt.65,104,106,107,430,431,443,721,799 Taken together, the evidence swine and avian influenza viruses also seem to become
at present suggests that, while H5N1 viruses can cause very apparent soon after exposure (e.g., within 5 days for most
serious illnesses in animals, milder cases are also possible. North American H3N2 swine influenza viruses and Asian
Mink seem to be susceptible to a variety of influenza lineage H5N1 HPAI viruses), although the incubation
viruses from other species,7,65,74,75,441,534,801 and two period for some H5N1 cases might be as long as 8-17
recently published studies suggest that H9N2 avian days.203,207,612,707,725,726,866,867
influenza viruses might be a concern on mink farms in
China.441,442 While morbidity rates in influenza virus- Clinical Signs
infected mink can approach 100%, mortality rates have
Seasonal human influenza
differed between outbreaks, and were probably influenced
by co-infections and other factors.7,65,74,75,441,534,801 During Uncomplicated infections with human influenza A
one extensive and severe outbreak caused by a avian viruses (including pandemic H1N1 virus) or influenza B
H10N4 virus, the morbidity rate was nearly 100% and the viruses are usually characterized by nonspecific symptoms
mortality rate was 3%.7,65 In contrast, an H9N2 avian and upper respiratory signs, which may include fever,
influenza virus and an H3N2 swine influenza virus caused chills, anorexia, headache, myalgia, weakness,
few or no deaths.75,441 photophobia, sneezing, rhinitis, sore throat and a cough. 7,9-
12,31
Intestinal signs (vomiting, nausea, diarrhea, abdominal
In seals, the case fatality rate was estimated to be 20% pain), otitis media and febrile seizures can also occur,
in one outbreak caused by an H7N7 virus, and 2-4% in an especially in children, and dehydration is a particular
outbreak caused by an H4N5 virus.7,562 Explosive concern in very young patients.9,10,30,31,868 Most people
epidemics in seals are thought to be exacerbated by high recover from the acute, uncomplicated illness within a
population densities and unseasonably warm temperatures, week, but coughing and tiredness may persist longer, and
as well as co-infections.65,763 secondary bacterial or viral infections can exacerbate or
Reports of illnesses caused by the 2009 pandemic prolong the symptoms.7,9,30,31
H1N1 viruses in pet cats, dogs, ferrets and zoo animals have More severe respiratory syndromes, including
been uncommon, but a number of these cases were severe pneumonia, are possible,9,11,12,30,31 and deterioration can
or fatal.67-73,510-514,517,518,520,522,523,527 In one outbreak at a cat occur rapidly in these cases.35,869 Severe primary viral
colony, half of the cats had clinical signs, and 25 of the 90 pneumonia and/or acute respiratory distress syndrome, as
cats died.521 However, it is possible that milder cases have well as multiple organ failure and other serious
not been recognized. Two surveys found increasing levels syndromes, occurred in a small percentage of cases during
of antibodies to pandemic H1N1 virus among cats, with the 2009-2010 pandemic, and affected an unusually large
reported rates as high as 22% and 31% among pet cats in number of children and young adults.32,38,44,47,865,869-871
the U.S. and China, respectively, and 11% among cats in Influenza can also result in the decompensation or
animal shelters in China.732,798 Infections with this virus exacerbation of serious underlying diseases such as
have also stimulated renewed research into the possibility

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 Influenza
chronic lung conditions, cardiac conditions, poorly were also reported in two adults.243 However, severe lower
controlled diabetes, chronic renal failure or end-stage liver respiratory disease, which developed into respiratory
disease.32,35,37,38,45,871-873 Other possible complications failure, occurred in an immunocompromised woman who
include various neurological syndromes including had serious underlying conditions.241
encephalitis, myositis (e.g., benign acute childhood
H7N9 LPAI viruses in China
myositis), rhabdomyolysis and myocarditis.11,30,139-141,868
Influenza-related deaths are usually the result of Most clinical cases caused by H7N9 viruses in China
pneumonia, the exacerbation of a cardiopulmonary have been serious.231,258,262,695,880-882 The most common
condition or other chronic disease, or complications symptoms were fever and coughing, but a significant
associated with age or pregnancy.31 number of patients also had dyspnea and/or hemoptysis on
initial examination, and most cases progressed rapidly to
Influenza C virus infections are mainly characterized severe pneumonia, frequently complicated by acute
by mild upper respiratory disease, with some studies also respiratory distress syndrome and multiorgan
reporting gastrointestinal signs or otitis; however, more dysfunction.696,866,883 Diarrhea and vomiting were
severe cases with lower respiratory signs including sometimes reported, but conjunctivitis was uncommon,
pneumonia have been reported.7,129-138,157 Fever was a and most patients did not have nasal congestion or
common symptom in some studies,132,134-136 but a study of rhinorrhea as the initial signs.866,884
young adults in Finland found that most had mild upper
respiratory signs without fever.138 Neurological signs have A few uncomplicated cases were characterized by
been reported in a few cases, and included seizures/ mild upper respiratory signs or fever alone, especially in
unconsciousness in an infant, and drowsiness and children.258,725,866,881,884,885 Some of these cases may have
hemiparesis in a child.136,874 been mild due to prompt treatment with oseltamivir, but
others were admitted to the hospital for observation alone
Avian influenza infections in humans or identified only after the person had
Asian lineage H5N1 HPAI viruses and reassortants recovered.258,866,881,885 At least one asymptomatic infection
has been reported in an adult.695,866 Antibodies to H7N9
Most infections with Asian lineage H5N1 HPAI
viruses in healthy poultry or live bird market workers
viruses have been severe.53,108,237 The initial signs are often
suggest that some mild cases or asymptomatic infections
a high fever and upper respiratory signs resembling human
might not have been diagnosed.609,882,886-888
seasonal influenza, but some patients may also have
mucosal bleeding, or gastrointestinal signs such as Other avian influenza viruses
diarrhea, vomiting and abdominal pain.203,207,875 Lower Mild illnesses, characterized by conjunctivitis and/or
respiratory signs tend to develop soon after the onset of the upper respiratory signs, have been reported in a number of
illness.203,207 Most patients deteriorate rapidly, and serious people infected with various H7 LPAI or HPAI viruses and
complications including multiorgan dysfunction are H10N7 viruses.172,561,598,614,615,618,722,724,889-893 One H7N7
common in the later stages.203,207,875 Milder cases have HPAI virus, which caused only mild illness in most people,
been reported occasionally, particularly among resulted in fatal acute respiratory distress syndrome and
children.237,876 Rapid treatment with antiviral drugs may other complications in one otherwise healthy person.722 His
have been a factor in some mild cases;877-879 however, at initial symptoms included a persistent high fever and
least one child with upper respiratory signs made an headache but no signs of respiratory disease. Severe
uncomplicated recovery after antibiotic treatment alone. 876 pneumonia was reported in a person infected with an LPAI
Three infections with Asian lineage H5N6 HPAI H7N2 virus who had serious underlying medical
viruses in older adults were severe, with fever and severe conditions.729 He was hospitalized but recovered. A 20-
respiratory signs in at least two patients.232-234 One of these year-old woman infected with an H6N1 virus in China had a
cases was fatal; the other patient required mechanical persistent high fever and cough, progressing to shortness of
ventilation but recovered after treatment with oseltamivir breath, with radiological evidence of lower respiratory tract
and antibiotics.233,234 (Details of the third case have not disease.616 She made an uneventful recovery after treatment
been published.) A child infected with an H5N6 virus had with oseltamivir and antibiotics. Three older adults with
a mild illness with prompt recovery.232-234 H10N8 infections in China developed severe lower
respiratory tract disease, progressing in some cases to
Eurasian lineage H9N2 LPAI viruses
multiple organ failure and septic shock, and two of these
Most illnesses caused by H9N2 viruses have been cases were fatal.617,730
reported in children and infants.108,235,237,238,241-244 These
cases were usually mild and very similar to human Swine influenza virus infections in humans
influenza, with upper respiratory signs, fever, and in some Most laboratory-confirmed, symptomatic swine
cases, gastrointestinal signs (mainly vomiting and influenza virus infections have been characterized by upper
abdominal pain) and mild dehydration.108,235,237,238,241-244 respiratory signs that resemble human influenza, including
All of these patients, including a 3-month-old infant with gastrointestinal signs in some patients, although acute
acute lymphoblastic lymphoma,241 made an uneventful parotitis was reported in a 6-year-old with H3N2 influenza,
recovery. Acute, influenza-like upper respiratory signs and one young patient had only fever and vomiting.7,14,98,99,

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 Influenza
286,293,498,584,586,591,593,595,597,702,706,707,894,895
In a recent series of dangerous human pathogens such as H5N1 HPAI
infections caused by North American triple reassortant viruses.53,207
H3N2 viruses, eye irritation appeared to be more common Resistance to antiviral drugs can be detected either
than with seasonal influenza viruses.707 Most healthy people with phenotypic tests or by gene-based testing to detect
infected with these H3N2 viruses had a mild illness, molecular markers of resistance.897 The need to perform
although young children were sometimes hospitalized for susceptibility testing depends on the composition of
dehydration.98,99,286,293,584,586,591,593, circulating viruses and the individual case.897 These tests
595,702,707,894
The illness was also mild and flu-like in one are available in a limited number of laboratories and take
cancer patient undergoing chemotherapy, who became several days to perform.897
infected with a European swine influenza virus.597 Because people have antibodies to the subtypes found
Serological evidence suggests that mild or asymptomatic in circulating influenza viruses, serology is not generally
cases might occur sporadically among people who are useful for the routine diagnosis of seasonal influenza.31
occupationally exposed.7,14,159,337,599-604 However, zoonotic influenza virus infections are
Swine influenza viruses of various subtypes have occasionally diagnosed retrospectively by serology. 703
occasionally caused pneumonia, serious illnesses and Although a rising titer must be seen for a definitive
deaths, usually in people who had underlying health diagnosis, single titers may be helpful in some
conditions or were immunocompromised by disease or circumstances. Tests used to detect influenza virus
pregnancy.99,286,585,588-590,595,596,702,707,896 A few serious or antibodies in humans include hemagglutination inhibition,
fatal cases occurred in healthy people. virus neutralization, enzyme immunoassays and
Equine and canine influenza virus infections in complement fixation.897,903 The microneutralization assay
humans is considered to be the most reliable test for detecting
antibodies to avian influenza viruses.207,237 People infected
There are no reports of clinical cases caused by
with some avian influenza viruses did not seroconvert,
natural exposure to equine influenza viruses or canine
even in virologically confirmed cases.614,724
influenza viruses, although human volunteers inoculated
with an equine influenza virus became ill.7 Infections caused by influenza C viruses
RT-PCR or culture can be used to diagnose influenza
Diagnostic Tests C.904,905. It can be difficult to isolate these viruses in cell
Infections caused by influenza A and B viruses lines, and although they can be isolated in embryonated
A number of assays, similar to those used in animals, eggs, this technique is not widely available in diagnostic
can diagnose influenza A and B infections in humans.31,897 laboratories.905
Upper respiratory samples are generally collected for
Treatment
routine seasonal influenza diagnosis, but samples from the
lower respiratory tract are appropriate in some cases. 53,207 Supportive care for uncomplicated influenza in
RT-PCR techniques are now the method of choice for humans includes fluids and rest. Adjunct and supportive
detecting and subtyping human influenza viruses in many treatments for severe, hospitalized cases vary, and can
laboratories, due to their speed and sensitivity.897,898 These include various drugs, including antibiotics to treat or
tests can also be used for zoonotic influenza virus prevent secondary bacterial pneumonia, and mechanical
infections.207,725,899,900 Virus isolation can be done, ventilation.32
although traditional techniques take 3-14 days, and are too Two groups of antiviral drugs – adamantanes
slow for the initial diagnosis and management of the (amantadine, rimantadine), and neuraminidase inhibitors
case.45,897 Some newer methods (e.g., shell vial-based (zanamivir, oseltamivir, peramivir and laninamivir) – are
culture techniques) are faster, if available.897 Antigen- used to treat some cases of influenza.9,11,30,31,45,906,907
detection assays used in humans include Adamantanes are active against human influenza A
immunofluorescence and immunoassays such as viruses, while neuraminidase inhibitors can be used in
ELISAs.9,31,45,898 Commercial rapid diagnostic test kits can both influenza A and influenza B
provide a diagnosis within 15 minutes, but are less infections.9,11,12,23,31,45,906,907 Antiviral drugs are most
sensitive than some other methods (e.g., RT-PCR), differ effective if they are started within the first 48 hours after
in complexity and in the viruses they can distinguish, and the clinical signs begin, although they may also be used in
may not detect novel infections including zoonotic severe or high risk cases first seen after this
viruses.31,45,53,897,898,901,902 Testing that identifies the time.11,12,23,31,45,906,907 Specific recommendations for
presence of influenza A, but does not detect the subtypes antiviral use can vary, but these drugs are usually
found in common human influenza viruses, might indicate recommended for severe cases of influenza, or infections
a novel, possibly zoonotic, influenza virus.53 Testing for that have an elevated risk of complications, and may also
novel influenza viruses is generally performed by state, be employed for some milder cases of seasonal
regional or national public health laboratories, and in some influenza.31,45,203,207,314,875,908 There has been some debate
cases by reference laboratories capable of handing about the benefits of oseltamivir for uncomplicated
seasonal influenza in healthy patients.9,30,45,906,909-913 The

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 Influenza
side effects of specific drugs vary, but can include groups, such as the elderly, is consistently
gastrointestinal and CNS effects.31,45 recommended.31,924
The development of antiviral resistance is a concern, Antiviral drugs may be used for prophylaxis in some
especially if drugs are used indiscriminately. Resistance high-risk populations such as the elderly or
can develop rapidly in influenza viruses, and may even immunocompromised, or people may be monitored and
emerge during treatment.7,9,30,31 At one time, adamantanes treated at the first sign of disease. 30,31,45,906 The use of
were used most often to treat seasonal influenza in the antiviral prophylaxis should be balanced against the risk
U.S.; however, many H1N1, H3N2, and influenza B of encouraging the emergence of drug-resistant strains.45
viruses had become resistant to these drugs by the 2006- Other preventive measures include avoiding close contact
2008 flu seasons.7,9,30,897,906,907 After antiviral drug with people who have influenza symptoms, and common
recommendations changed, seasonal H1N1 influenza sense hygiene measures such as frequent hand washing
viruses rapidly became resistant to oseltamivir, although and avoidance of unnecessary hand contact with the eyes,
many lost their resistance to adamantanes.897 These H1N1 nose or mouth.31,53,97,925 To protect others, the mouth and
viruses co-circulated with adamantane-resistant, nose should be covered when coughing or
oseltamivir-sensitive H3N2 and influenza B viruses, sneezing.31,97,925 Recommendations on the use of face
complicating treatment decisions.897 This pattern changed masks, respirators, gloves and other barrier precautions
with the introduction of pandemic H1N1, and during the vary, and current setting-specific guidelines (e.g., for
2015-2016 influenza season, many of the circulating hospitals vs. the community) should be consulted. 31,32,673
seasonal influenza viruses in the U.S., including pandemic The effectiveness of face masks and respirators in
H1N1, were resistant to adamantanes and sensitive to decreasing influenza virus transmission is still under
neuraminidase inhibitors.31 There have also been reports of investigation, although some studies suggest that they
seasonal influenza A viruses resistant to both drug classes, may reduce the amount of virus transmitted by the
and oseltamivir-resistant influenza B viruses have been wearer, and/or provide some protection to the
found.897,914 wearer.672,673,675,926-930
Asian lineage H5N1 HPAI viruses and Chinese H7N9 Additional measures that have been recommended
LPAI viruses are usually sensitive to oseltamivir and during pandemics or outbreaks caused by novel viruses
resistant to adamantanes, at present, although this could include avoidance of crowds and gatherings, cancellation
change.53,207,259,640,883,915,916 One recent study documented of social events, and voluntary self-isolation of individuals
pre-existing resistance to neuraminidase inhibitors, at low who develop influenza-like signs (with the exception of
levels, among avian influenza viruses in wild birds, and in necessities such as seeking medical care).925,931-936
9% of viruses isolated from swine that contain the N2
Zoonotic influenza viruses
neuraminidase (H1N2, H3N2 and H9N2).917 Another
study found a high prevalence of amantadine resistance in Protective measures for zoonotic influenza viruses
some lineages of swine influenza viruses in North include controlling the source of the virus (e.g.,
America.918 Published reports suggest that neuraminidase eradicating viruses from domesticated birds, closing
resistance is currently uncommon among swine influenza infected poultry markets); using sanitation and hygiene
viruses in the U.S. and Germany.918,919 measures such as hand washing; avoiding contact with
sick animals or animals known to be infected; and
Guidance on the influenza viruses circulating during
employing personal protective equipment where
the current season, with treatment recommendations, is
appropriate (for instance, when working with infected
often available from national or local health authorities
birds or swine).53,97,203,704 While zoonotic infections are
(e.g., the CDC in the U.S.). Antiviral susceptibility testing
usually acquired during close contact with animals, 53,97
can be done, but it is too slow to guide the initial
aerosolized viruses may be present in confined areas such
treatment, which should be started during the period of
as production barns, where large numbers of swine are
maximum virus susceptibility.897
concentrated.676,677 Because HPAI viruses have been found
Prevention in meat and/or eggs from several avian species,164,418,660-
666,778,937-941
Annual vaccines, usually given in the fall before the careful food handling practices are important
flu season (or as appropriate for local patterns of virus when working with raw poultry or wild game bird
circulation), are available for influenza A and B. 7,9,268,920,921 products in endemic areas, and all poultry products should
They contain the viral strains considered most likely to be completely cooked before eating.53,826,942 Swine
produce epidemics during the following winter, including influenza viruses can also be inactivated by
pandemic H1N1 virus, and are updated annually. Details cooking,97,943,944 although these viruses are respiratory
on vaccine efficacy, vaccine types, and current pathogens and are not likely to be present in retail meat.642
recommendations are available from government sites More detailed recommendations for specific groups
(e.g., the CDC in the U.S.) and professional advisory at risk of exposure, including the general public, have
groups.31,276,922,923 Immunization recommendations may been published by some national agencies (e.g., the CDC,
differ between countries, although vaccination of some the Department of the Interior and U.S. Geological
Survey National Wildlife Health Center in the

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 Influenza
U.S.,)53,97,826,945 and international agencies such as the 2009-2010 pandemic
World Health Organization. In some cases, Serological studies have estimated that approximately
recommendations may include antiviral prophylaxis 30-50% of all school-aged children, and a smaller
and/or vaccination for human influenza to reduce the risk percentage of the entire population (10-40% worldwide)
of reassortment between human and animal influenza were infected during the initial stages of the 2009-2010
viruses.53 Currently, the CDC recommends that people pandemic.37,951,952 Overall, the pandemic H1N1 virus has
who are at elevated risk of complications with human caused relatively mild illnesses, and the estimated case
influenza viruses avoid pigs and pig barns when fatality rate for this virus is less than 0.5%, with a number
attending fairs in North America, and that other people of estimates suggesting that it is less than 0.05%.37,47
take precautions to reduce the risk of infection with Nevertheless, an elevated number of patients developed
swine influenza viruses. 97 When visiting a physician for viral pneumonia during this pandemic, and case fatality
an illness that began soon after contact with animals, the rates in younger age groups were higher than with
potential for zoonotic exposure should be mentioned. seasonal influenza viruses.35,37,44,45,869,953 Most hospitalized
or severely affected patients were children and young
Morbidity and Mortality adults, with relatively few patients older than 50 years and
Human influenza A and B viruses even smaller numbers older than 60.36,37 The relative
Human influenza can occur as a localized outbreak, sparing of older populations appeared to result from
epidemic, or pandemic, or as sporadic cases. 8 Historical immunity to similar, previously circulating viruses
evidence suggests that pandemics occur every 10 to 40 (pandemic H1N1 was antigenically very similar to the
years.7,11,14 Epidemics are seasonal in temperate regions, 1918 virus), and possibly other factors.33,36,47,954-957 The
typically beginning after school starts in the fall, and concentration of severe illnesses mainly in younger,
spreading from children to adults, although some virus healthier age groups is thought to have contributed
transmission occurs outside this time. 7,9,946 In tropical and significantly to the low overall mortality rate, and many
subtropical areas, influenza patterns are very diverse, seriously ill patients recovered with hospitalization and
with transmission occurring year-round in some intensive care.32,37 However, older patients who became
countries, and seasonal epidemics, sometimes coinciding infected had an elevated risk of severe illness and death.47
with the rainy season or occurring in two peaks, in The prevalence of pre-existing conditions among
others.920,921,947-949 seriously ill children differed between studies, but
Uncomplicated infections with seasonal influenza predisposing conditions (e.g., asthma, immunosuppression,
viruses are rarely fatal in most healthy people, although neurological diseases) were relatively common in some
the morbidity rate can be high.7-12 Approximately a third series.32,958,959 Nevertheless, a significant number of serious
of influenza virus infections are thought to be or fatal cases were reported in healthy children or young
asymptomatic.47 Groups at higher risk for severe illness adults.32,35,37,38,44,45,47,869,960,961 Obesity and pregnancy were
include the elderly; young children (due to risks from recognized as risk factors for more serious illness during
complications such as severe dehydration); people with this pandemic.31,43 The impact of this virus was also greater
chronic respiratory or cardiovascular disease and various in indigenous groups.33,37,47 The reason is still uncertain, but
other medical conditions; members of some ethnic groups might involve access to healthcare, concurrent illnesses,
at high risk (see pandemic H1N1, below); and those who increased crowding or other factors.37,47
are immunosuppressed, including pregnant women.9,11,12,30- Influenza C
38,43
Obesity was first recognized as a risk factor during the
Serological studies suggest that many people are
2009-2010 pandemic.31,43
exposed to influenza C viruses in childhood, although
Since 1968, H3N2 influenza A viruses have caused infections can continue to occur in adults.136-138,962-965 One
the most serious epidemics with the highest mortality recent study from Scotland found two peaks of mild
rates.30,31 Except after the introduction of a new virus, over influenza C-related illness, one in children and a second in
90% of influenza-related deaths occur in the elderly.30,268 adults over the age of 45 years.157 At one time, these
Morbidity and mortality rates usually increase during viruses were thought to cause only sporadic cases of
influenza A pandemics, sometimes dramatically.7,9,14,31,46,53 influenza and minor localized outbreaks.7,9,129 However, in
The pandemic of 1918 is notorious for its severity, with 2004, a nationwide influenza C epidemic was reported in
some estimates suggesting a morbidity rate of 25-40% and Japan.904 Infections seem to be most serious in very young
case fatality rate of 2-5%.8 It should be noted that antiviral children. In one study, 30% of children hospitalized with
drugs and antibiotics were not available at the time, and severe influenza C infections were less than two years old,
intensive care procedures were less effective. After a and an additional 12% were between the ages of 2 and 5
pandemic, an influenza virus (or its variants950) usually years.132
becomes established in the population and circulates for
years.7,8,14,124 Influenza B viruses can cause epidemics, but Zoonotic swine influenza
they have not, to date, been responsible for pandemics. 7 The overall prevalence of swine influenza virus
infections in humans is uncertain. While cross-reactivity to

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 Influenza
human influenza viruses may be an issue, serological Asian lineage H5N6 viruses have caused a few severe
studies suggest that some people who work with pigs have clinical cases in patients aged 49 years or older; however,
been exposed to swine influenza viruses.7,14,159,309,337,599- one infection in a child was mild.232-234
604,606,607,609,610,966,967
If most infections resemble human
H7N9 avian influenza viruses in China
influenza, they may not be investigated and recognized as
zoonoses. Virologically confirmed clinical cases caused by Approximately 680 laboratory-confirmed clinical
H1N1, H1N2 and H3N2 viruses have been reported cases, with at least 275 fatalities, have been caused by
sporadically since the 1970s (with one localized outbreak H7N9 LPAI viruses in China, as of September 2015. 231
in 1976), and more regularly in recent years. 7,14,97- They mainly occurred in three waves to date, the first
99,159,286,287,293,584-597,702,706,707,894-896
Although zoonotic cases between February and May 2013, the second from
have also been seen in Europe and Asia, most recent cases October 2013 to May 2014, and the third beginning in
were documented in the U.S., where this disease has been Fall 2015, with sporadic cases reported between
reportable since 2005.97 Approximately one case was outbreaks.258,259,261,262,888,978 These viruses are circulating
documented every 1-2 years in the U.S. at one time; subclinically among poultry in China, and human cases
however, this increased to 21 cases between 2005 and June have mainly been associated with live bird poultry
2011, 13 cases from August 2011 to April 2012, and 306 markets; however, infected farms have also resulted in at
confirmed cases (mainly associated with fairs) in summer least one human illness. 263,264,612,703,704,888,979,980 Most
2012.97-99 This increase may be related to changes in swine reported cases have been serious, except in children, who
influenza viruses (particularly the establishment of triple often (though not always) presented with mild
reassortment H3N2 viruses in swine populations, and their illnesses.258,263,264,695,866,880,881 Elderly people were
reassortment with 2009 pandemic H1N1 virus), but other overrepresented among the clinical cases, probably due
factors, such as increased surveillance and new reporting to increased exposure and/or increased
requirements may also play a role.96-99 susceptibility.612,880,981 As of October 2014, case fatality
rates were approximately 36% to 48% in hospitalized,
Many cases of swine influenza (including most of the
laboratory confirmed cases during the first two
recently reported cases in the U.S.) have been seen in
waves,258,264,882,978 with the risk of death among
children, but adults are also affected.98,99,286,293,584-586,588-
591,593,595,596,702,707,894,896 hospitalized patients increasing significantly with
While most cases have been mild and
age.258,882 Concurrent diseases or predisposing causes
resembled human influenza, a few severe or fatal illnesses
have been reported in a significant number of patients,
have also been reported, often but not always in people who
although serious cases and fatalities also occurred in
had underlying health conditions or predisposing
previously healthy individuals. 258,264,612,641,695,725,866,880,982
factors.98,99,286,293,584-586,588-591,593,595,596,
702,707,894,896 The likelihood of additional, undiagnosed mild or
asymptomatic infections is still being assessed, although
Asian lineage H5 avian influenza viruses few cases were detected during national virological
Between 1997 and February 2016, Asian lineage sampling of people with influenza-like illnesses.258,264,881
H5N1 viruses were responsible for more than 800 Some initial serological studies reported no H7N9
laboratory-confirmed human infections, generally as the reactivity among poultry market workers, healthcare staff,
result of close contact with poultry231 Most patients were patient contacts and other populations.259,726,983-986
young and had no predisposing conditions.207 The case However, several surveys have now detected antibody
fatality rate for all laboratory confirmed cases reported to titers to H7N9 viruses in up to 17% of poultry workers or
WHO has consistently been about 59-60% in the last few live bird market workers, with two studies documenting
years.231,611,968 However, it differs between countries, and recent increases in seroprevalence.609,886-888 These studies
is particularly low in Egypt, where 28% of confirmed, report that seroprevalence rates are low (≤ 1%) in the
suspect and probable cases between 2006 and 2010 were general population,886,887 with one survey also
fatal.875,877,879,969-971 A high proportion of the reported cases documenting low seroprevalence (2%) in veterinarians. 609
in Egypt occurred in young children, and their young age, Taking into account the serological studies, some authors
early diagnosis and, treatment-related factors, as well as have speculated that the overall case fatality rate might be
the virulence of the circulating viruses, might be factors in as low as <1% to 3%, if milder cases are also accounted
the relatively high survival rate.877-879 Antibodies to H5 for.258,882
viruses have been reported (generally at low rates of
Eurasian lineage H9N2 avian influenza viruses
seroconversion) in some poultry-exposed populations that
have no history of severe H5N1 disease, fueling Clinical cases caused by H9N2 viruses have mainly
speculation on the likelihood of asymptomatic or mild been reported in children.53,108,231,235-238,242-244 Most cases,
infections.431,609,620,621,623,629,972-976 Rare, laboratory including an infection in an immunocompromised infant,
confirmed, asymptomatic or mild cases have also been have been mild, and were followed by uneventful
recognized.237,876,977 Recent prospective studies recovery. Severe illness was reported in an adult with
documented seroconversion in rare instances, but detected serious underlying medical conditions.241 Serological
no clinical cases.619,621,622 studies suggest that exposure to H9N2 viruses may occur

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 Influenza
in some people who are exposed repeatedly to poultry in United States Department of Agriculture (USDA) Animal
endemic areas,236,239,240,605,609,620,621,623,624,976,987-989 and a and Plant Health Inspection Service (APHIS).
prospective study of adults with poultry exposure in rural https://www.aphis.usda.gov/aphis/home/
Thailand reported rare instances of seroconversion,
USDA APHIS. Biosecurity for the Birds
although no clinical cases were detected.619
https://www.aphis.usda.gov/aphis/ourfocus/animalhealth/a
Other avian influenza viruses nimal-disease-information/avian-influenza-
With the exception of the H7N9 viruses in China, disease/birdbiosecurity
most reported infections with H7 viruses in healthy people United States Geological Survey (USGS). National
have been mild, whether they were caused by an LPAI or Wildlife Health Center. List of Species Affected by H5N1
HPAI virus; however, one H7N7 HPAI virus caused a (Avian Influenza)
fatal illness in a healthy person, while affecting others only http://www.nwhc.usgs.gov/disease_information/avian_infl
mildly.172,561,615,722,724,889-893 Mild signs were reported in uenza/affected_species_chart.jsp
poultry workers infected with an H10N7 virus, 614 but
H10N8 viruses caused fatal infections in two elderly USGS National Wildlife Health Center. Wildlife Health
patients in China and a serious illness in a 55-year- Bulletin #05-03 (with recommendations for field
old,617,730 and a young woman infected with an H6N1 virus biologists, hunters and others regarding contact with wild
in China developed lower respiratory tract birds
complications.616 The possibility of other, unrecognized http://www.nwhc.usgs.gov/publications/wildlife_health_b
infections may be suggested by the occurrence of ulletins/WHB_05_03.jsp
antibodies, generally at a low prevalence, to H4, H6, H7, Department of the Interior. Appendix H: Employee Health
H10, H11 and H12 viruses (as well as H5 and H9 viruses) and Safety Guidance for Avian Influenza Surveillance and
in people who are exposed to poultry or Control Activities in Wild Bird Populations
waterfowl.236,237,239,240,600,619,621,625-629,631,988,990-992 http://www.doi.gov/emergency/pandemicflu/appendix-
h.cfm
Internet Resources
World Health Organization. Zoonotic Influenza
Centers for Disease Control and Prevention (CDC). http://www.who.int/influenza/human_animal_interface/en/
Avian Influenza
World Organization for Animal Health (OIE)
http://www.cdc.gov/flu/avianflu/
http://www.oie.int
CDC. Seasonal Influenza . (with links to avian, swine and
OIE Manual of Diagnostic Tests and Vaccines for
other influenza viruses)
Terrestrial Animals
http://www.cdc.gov/flu/
http://www.oie.int/international-standard-
Department of the Interior. Appendix H: Employee Health setting/terrestrial-manual/access-online/
and Safety Guidance for Avian Influenza Surveillance and
OIE Terrestrial Animal Health Code
Control Activities in Wild Bird Populations
http://www.oie.int/international-standard-setting/terrestrial-
http://www.doi.gov/emergency/pandemicflu/appendix-
code/access-online/
h.cfm
Prevention and Control of Influenza. Recommendations of Acknowledgements
the Advisory Committee on Immunization Practices
http://www.cdc.gov/vaccines/hcp/acip-recs/vacc- This factsheet was written by Anna Rovid Spickler, DVM,
specific/flu.html PhD, Veterinary Specialist from the Center for Food
Security and Public Health. The U.S. Department of
Public Health Agency of Canada (PHAC). Influenza Agriculture Animal and Plant Health Inspection Service
http://healthycanadians.gc.ca/diseases-conditions- (USDA APHIS) provided funding for this factsheet
maladies-affections/disease-maladie/flu-grippe/index- through a series of cooperative agreements related to the
eng.php development of resources for initial accreditation training.
PHAC. Pathogen Safety Data Sheets The following format can be used to cite this factsheet.
http://www.phac-aspc.gc.ca/lab-bio/res/psds-ftss/index- Spickler, Anna Rovid. 2016. Influenza. Retrieved from
eng.php http://www.cfsph.iastate.edu/DiseaseInfo/factsheets.php.
The Merck Manual
http://www.merckmanuals.com/professional
The Merck Veterinary Manual
http://www.merckvetmanual.com/

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 Influenza
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