PNAS - Emergence and predominance of an H5N1 influenza variant

[sharon sanders]

Emergence and predominance of an H5N1 influenza
variant in China
G. J. D. Smith?, X. H. Fan?, J. Wang?, K. S. Li?, K. Qin?, J. X. Zhang?, D. Vijaykrishna?, C. L. Cheung?, K. Huang?,
J. M. Rayner?, J. S. M. Peiris?, H. Chen?, R. G. Webster???, and Y. Guan??
?State Key Laboratory of Emerging Infectious Diseases, Department of Microbiology, Li Ka Shing Faculty of Medicine, University of Hong Kong,
21 Sassoon Road, Pokfulam, Hong Kong Special Administration Region, China; and ?Virology Division, Department of Infectious Diseases, St. Jude Children?s Research Hospital, Memphis, TN 38105

Contributed by R. G. Webster, September 20, 2006

The development of highly pathogenic avian H5N1 influenza
viruses in poultry in Eurasia accompanied with the increase in
human infection in 2006 suggests that the virus has not been
effectively contained and that the pandemic threat persists. Updated
virological and epidemiological findings from our market
surveillance in southern China demonstrate that H5N1 influenza
viruses continued to be panzootic in different types of poultry.
Genetic and antigenic analyses revealed the emergence and predominance
of a previously uncharacterized H5N1 virus sublineage
(Fujian-like) in poultry since late 2005. Viruses from this sublineage
gradually replaced those multiple regional distinct sublineages and
caused recent human infection in China. These viruses have already
transmitted to Hong Kong, Laos, Malaysia, and Thailand, resulting
in a new transmission and outbreak wave in Southeast Asia.
Serological studies suggest that H5N1 seroconversion in market
poultry is low and that vaccination may have facilitated the
selection of the Fujian-like sublineage. The predominance of this
virus over a large geographical region within a short period directly
challenges current disease control measures.
influenza A
molecular epidemiology
virus evolution
Extensive surveillance and genetic studies have revealed that
highly pathogenic avian influenza H5N1 viruses have become
first predominant and then endemic in poultry in southern
China and Southeast Asia since 2003 (1). This endemicity
resulted in the establishment of multiple distinct regional sublineages
(2). The recognition of multiple different H5N1 sublineages
makes it possible to identify the source and to understand
the evolutionary and transmission pathways of H5N1
viruses that have become widespread in Southeast Asia, Europe,
and Africa.
Since the H5N1 influenza virus caused the first outbreak in
migratory waterfowls at Qinghai Lake in May 2005 (3), a new
transmission and outbreak wave was initiated. The virus expanded
its geographical distribution and caused outbreaks in poultry in over
30 countries from Central Asia, the Middle East, Europe, and
Africa (4). This expansion led directly to a marked increase in
human infection cases and escalated the pandemic threat. In the
first 8 months of 2006, the World Health Organization confirmed
96 cases from 9 countries; whereas for the whole of 2005 there was
a total of 95 cases from 5 countries (5). In addition, in Indonesia
recently there were suspected cases of human-to-human transmission
involving members of an extended family, and the infection
sources of other human cases have not been identified (6).
In China, despite a compulsory program for the vaccination of all
poultry commencing in September 2005 (7), H5N1 influenza virus
has caused outbreaks in poultry in 12 provinces from October 2005
to August 2006 (4). At the same time, 22 human infection cases have
been confirmed from 14 provinces since November 2005 (4, 5).
Some of those cases were residents of metropolitan areas remote
from poultry farms, such as Guangzhou, Wuhan, and Shanghai (4).
Furthermore, there were no obvious poultry outbreaks reported in
neighboring markets or farms before or after the onset of those

 

[sharon sanders]

Re: Emergence and predominance of an H5N1 influenza

Re: Emergence and predominance of an H5N1 influenza

human infections. Therefore, whether those people were infected
locally and directly from affected poultry or other sources, including
humans, is still unknown. This situation directly challenges current
pandemic preparedness plans, raising concern that a pandemic
could emerge not only from the countryside but also from an urban
area, just as severe acute respiratory syndrome emerged from the
live-animal markets of Guangzhou and the Pearl River delta (8, 9).
Here we report updated virological and epidemiological findings
from our market surveillance in southern China. Epidemiological
analysis showed that H5N1 influenza viruses were
continued to be perpetuated in poultry in each of the provinces
tested, mainly in domestic duck and geese. Genetic analysis
revealed that an H5N1 influenza variant had emerged and
become predominant in each of the provinces, replacing those
previously established multiple sublineages in different regions
of southern China. This virus had also transmitted to Hong
Kong, Laos, Malaysia, and Thailand. Serological studies suggested
that the seroconversion rate in poultry in China is low and
that the emergence and predominance of this H5N1 strain may
be associated with vaccination in poultry. Genetic findings also
revealed that these viruses also were responsible for all recently
reported human infection cases in China. The predominance of
this virus over a large geographical region within a short period
questions the efficacy of current disease control measures in
poultry and revealed that a new transmission and outbreak wave
has been initiated from China to Southeast Asia since early 2006.
Results
Surveillance. From July 2005 to June 2006 our influenza surveillance
in live-poultry markets in six provinces of southern China
showed that 1,294 of 53,220 (overall isolation rate 2.4%) poultry
were H5N1-positive (Fig. 1; see also Fig. 5 and Table 3, which are
published as supporting information on the PNAS web site). The
main body of H5N1 isolates was from duck and goose, with only
a small number isolated from chicken (chicken 0.5%, duck 3.3%,
goose 3.5%). The prevalence of H5N1 viruses in southern China
has increased when compared with the period July 2004 to June
2005 (overall, 0.9%; chicken, 0.2%; duck, 1.3%; and goose,
2.0%) (Fig. 1 and Table 3). A winter-seasonal peak was observed
from October 2005 to March 2006 as in previous years (1, 2),
during which H5N1 influenza viruses were isolated in each

 

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Re: Emergence and predominance of an H5N1 influenza

Re: Emergence and predominance of an H5N1 influenza

province tested (Fig. 1). However, an extension of the peak
season was observed in April to June 2006 because isolation rates
remained high in these warmer months (Fig. 1) (1, 2).
Comparison between different types of poultry shows that H5N1
viruses were mainly isolated from domestic duck and goose wherein
the viruses were prevalent year-round, whereas chicken tested
positive mostly during the winter (Fig. 1 and Table 3). It is notable
that H5N1-positive chicken were detected in 11 of the last 12
months, a marked increase from only 4 positive months in 2004
2005 (Fig. 1A). These findings indicate an escalation of H5N1
activity in poultry in 20052006 compared with previous years and
suggest that H5N1 influenza viruses have not been effectively
contained in this region and have maintained endemicity broadly in
poultry, especially domestic duck and goose.
Regarding virus names and sublineages, the following nomenclature
applies: BH goose, bar-headed goose; Ck, chicken; FJ,
Fujian; Dk, duck; GD, Guangdong; Gf, Guinea fowl; Gs, goose;
GX, Guangxi; GY, Guiyang; HK, Hong Kong; HN, Hunan; IDN,
Indonesia; Mixed, southern China isolates; QH, Qinghai; ST,
Shantou; YN, Yunnan; VNM, Vietnam; VNM2, second Vietnam
introduction in March 2005;. The year the virus was
identified is represented by the last two digits of the year: e.g.,
02, 2002.
Antigenic Analysis. Antigenic analysis with World Health Organization
H5N1 reference antisera with representative viruses from
different sublineages (see below) demonstrated a diversity of
reaction patterns that generally corresponded to their phylogenetic
relationships (Fig. 2; see also Table 4, which is published as
supporting information on the PNAS web site). All tested FJ-like
H5N1 viruses showed high HA inhibition (HI) titers to antiserum
ofAnhui106, a virus from the same sublineage. Those viruses had
moderate to low reactivity to antisera of IDN35706, Iraq106,
DkHN10104, and BH gooseQH1A05 but no reactivity with
antisera of IDN505, VNM120304, Turkey6559606, and
Whooping swanMongolia24405 (Table 4).
Numerical analysis of HI titers conducted to visualize similarity
between the antigenic reactivity of different viruses
showed that those FJ-like H5N1 viruses had a distinguishable
antigenic reaction pattern. However, three viruses from this
sublineage (CkGX46306, DkHN85606, and GsST
1844205) differed slightly (Fig. 2). This analysis also revealed
two other major reactivity groups, one of Indonesia isolates and
another that contained viruses from three different sublineages
(GD06, QH-like, and MixedVNM2) (Fig. 2). Four viruses
(VNM120306, CkGY357005, GsYN553905, and Gs
GY33706) representing four other distinct H5N1 sublineages
had different reactivity patterns and did not group with other
viruses (Fig. 2 and Table 4).
Phylogenetic Analysis. To better understand the increased prevalence
of H5N1 in poultry and the emergence of human infection in
China, 390 (30% of total new isolates) of those avian H5N1
influenza viruses isolated from July 2005 to June 2006 plus 16
viruses isolated from smuggled poultry and dead wild birds in Hong
Kong in early 2006 were sequenced and analyzed together with
sequences available from public databases. Phylogenetic analysis of
the HA gene revealed that 266 of 390 (68%) of those recent H5N1
viruses from southern China formed a previously uncharacterized
and distinct H5N1 sublineage (FJ-like) (Fig. 3 and 4A). Twentyeight
viruses isolated in Guiyang from November 2005 to January
2006 formed a sublineage (GY2) that is the sister group to the
FJ-like sublineage (Figs. 3 and 4A).Another three sublineages from
Guangdong (GD06, n
6), Guiyang (GY1, n
14), and Yunnan
(YN2, n
13) also were identified. A further 59 viruses grouped
in the MixedVNM2 sublineage, and only a single virus (GfST
134106) belonged to the QH-like sublineage currently circulating
in Africa and Europe (Fig. 3 and 4A). The remaining viruses
analyzed in this study belonged to previously reported sublineages
from China and Southeast Asia, except two isolates from Hunan
(CkHN224606 and CkHN229206), isolated in May 2006 that
do not fall with any of these sublineages (Fig. 4A).
The prototype virus of the FJ-like sublineage (DkFJ173405)
was detected in March 2005. From July to September 2005, only a
single strain of 33 sequenced viruses was FJ-like (Table 1). Remarkably,
from October 2005 onwards the percentage of FJ-like
viruses detected increased dramatically, until from April to June
2006, 103 of the 108 H5N1 poultry isolates tested (95%) were
FJ-like (Table 1). Viruses from other sublineages (YN2, GY2, GY1,
GD06, and MixedVNM2) were not detected in our surveillance
since October 2005, November 2005, March 2006, April 2006, and
May 2006, respectively. These findings reflect the process of FJ-like
viruses gradually becoming predominant in this region.
Phylogenetic analysis also revealed that the HA gene of five
recent human H5N1 viruses from different provinces of China (4,
10, 11) belong to this FJ-like sublineage and were most closely
related to poultry isolates (Fig. 4A). This finding suggests that
H5N1 human infection from China since November 2005 was
Fig. 1. Comparison of H5N1 influenza virus isolation rate (%) in chicken (A),
duck (B) and goose (C) from southern China, July 2004 to June 2006. Surveillance
was conducted in live-poultry markets in Fujian, Guangdong, Guangxi,
Guiyang, Hunan, and Yunnan.
Smith et al.

 

[sharon sanders]

Re: Emergence and predominance of an H5N1 influenza

Re: Emergence and predominance of an H5N1 influenza

directly from affected poultry. Furthermore, H5N1 viruses isolated
in early 2006 from neighboring regions of mainland China, including
the 16 Hong Kong viruses and two poultry isolates from Laos
and Malaysia, all joined the FJ-like sublineage (Fig. 4A).
Of those genotyped viruses, 99 of 137 (72%) were from this
predominant FJ-like sublineage, all of which belonged to H5N1
genotype Z, except for a single genotypeGvirus (GsGY179406)
(Fig. 4B) (1, 2). Thirty-two viruses from other sublineages belonged
to genotypes G and Z, whereas six GD06 viruses belonged to
genotype X, which has only previously detected in Hong Kong and
Guangdong (Fig. 4B) (1). Of the 16 H5N1 virus isolates from Hong
Kong in 2006, there were 7 genotype Z viruses, whereas 9 genotype
G viruses were isolated from dead wild birds, suggesting a different
ecology. Genotype G has seven gene segments in common with
genotype Z but has a GsGD-like PB2 gene (Fig. 4B) (1, 2). These
findings show that FJ-like H5N1 influenza viruses have become
predominant and panzootic in southern China since mid-October
2005, have gradually replaced other H5N1 sublineages, and have
been transmitted to Laos and Malaysia.
Molecular Characterization. All viruses characterized in this study
maintained the motif of multiple basic amino acids at the HA
cleavage site characteristic of highly pathogenic avian influenza.
Furthermore, these viruses also kill embryonated eggs within
24 h of inoculation, which also is a characteristic of highly
pathogenic avian influenza. However, those viruses from the
FJ-like sublineage have a Gln-Leu substitution at position -9
from the cleavage site (LRERRR-KR/G .The receptor-binding
pocket of HA1 retains amino acid residues Gln-222 and Gly-224
(H5 numbering used throughout) that preferentially bind to
a
2,3 -NeuAcGal linkages of avian cell-surface receptors (12, 13).
Other amino acid residues relevant to receptor binding were
identical to those of Gs/Gd -like viruses (14), with the exception
of Dk/HN/5128/05, which had Val-131-Met substitution.
Only six viruses from the FJ-like lineage (Ck/ST/3840/06
CkST/3923/06, Ck/FJ/11933/05, CkFJ1223905, Ck/FJ/
58406, and DkGX155006) had the Ser-31-Asn mutation in

 

[sharon sanders]

Re: Emergence and predominance of an H5N1 influenza

Re: Emergence and predominance of an H5N1 influenza

DkFJ173405 (Table 2; see also Table 5, which is published as
supporting information on the PNAS web site). Almost all sera
had 2- to 4-fold higher titers to CkGY357005 and DkYN
440005, which are from the GY2 and MixedVNM2 sublineages,
in comparison with DkFJ173405 (Tables 4 and 5).
Only four sera were neutralizing-negative against all three
representative strains, although they were positive in the HI
assay (Table 5). These findings suggest that chicken in southern
China are poorly immunized against FJ-like viruses in comparison
with other sublineages.
Discussion
The highly pathogenic H5N1 influenza virus currently panzootic
in Eurasian and African poultry populations is considered the
most likely candidate for a new pandemic influenza. The development
of more and more avian-to-human interspecies transmission
events in the last 12 months seems to favor such a
hypothesis (4, 5). Our results demonstrate that the emergence
and predominance of a H5N1 influenza virus sublineage in
China has initiated a new transmission wave in Southeast Asia.
The emergence of this FJ-like sublineage has had similar consequences
to the first wave of virus transmission throughout Southeast
Asia in early 2004 (1) and the second wave to Europe and
Africa that followed the Qinghai Lake H5N1 outbreak (3, 4). The
findings of our study show that this virus has replaced most of those
previously established regional sublineages across a large geographical
area in China (2). The predominance of this FJ-like virus
appears to be responsible for the increased prevalence of H5N1 in
poultry since October 2005 and recent human infection cases in
China (4, 5). Furthermore, it has already caused poultry outbreaks
in Laos, Malaysia, and Thailand and human disease in Thailand (4,
5). As such, it is likely that this variant has already initiated a third
wave of transmission throughout Southeast Asia and may spread
further in Eurasia. It is also probable that this virus will continue to
evolve to form other regionally distinct sublineages, as witnessed
with the H5N1 genotype viruses in the first and second transmission
waves (1, 2, 17).
The mechanism for the emergence and prevalence of FJ-like
H5N1 variant is still unknown. The compulsory vaccination of all
poultry was ordered in China beginning September 2005 (7), but
our data indicate that seroconversion rates are still low and that
poultry are poorly immunized against FJ-like viruses, which suggests
that the poultry vaccine currently used in China may only
generate very low neutralizing antibodies to FJ-like viruses in
comparison to other previously cocirculating H5N1 sublineages.
This situation could have helped to select for the FJ-like sublineage
in poultry, because our results also show that these viruses had
replaced the GY2 and YN2 virus sublineages, both of which had
high titers in the serological tests. As such, this information suggests
that the predominance of FJ-like viruses may be associated with
immune escape from the current vaccine strain in poultry.
Previously, we described the establishment of multiple sublineages
of H5N1 virus in southern China and Southeast Asia (2). The
emergence and replacement of these sublineages by FJ-like viruses

 

[sharon sanders]

Re: Emergence and predominance of an H5N1 influenza

Re: Emergence and predominance of an H5N1 influenza

within a short period highlights the difficulties faced in controlling
H5N1 virus in China. A complex ecology and highly diverse virus
populations make it almost impossible to capture each circulating
virus sublineage, evenwith the application of mass vaccination. This
complexity has resulted in recurrent H5N1 outbreaks in poultry in
different regions and has led to occasional human infection.
Since November 2005, 22 H5N1 human infection cases from 14
provinces of China have been reported (4, 5). It is noteworthy that
four of the provinces (Fujian, Guangdong, Shanghai, and Zhejiang)
with human cases have not recorded any outbreaks in poultry (4).

However, phylogenetic and antigenic analyses in this study clearly
show that those recent human H5N1 isolates from different provinces
are FJ-like viruses, which suggests that this virus may be
prevalent in an area much larger than we have identified. Given the
lack of systematic influenza surveillance in poultry at a national
level, the timely identification of the source of human infection is
almost impossible. Therefore, to understand and identify possible
infection sources and to avert a potential pandemic, comprehensive
influenza surveillance in both human and animal populations is
urgently required in H5N1-affected regions.

The repeated emergence of H5N1 variants from southern
China and their subsequent spread to other parts of the world
(1?3) makes it increasingly apparent that implementation of
effective control measures in this region is of paramount importance.
Such a system of control measures could be achieved
by integrated real-time virological and genetic information with
rapid diagnostic approaches and vaccine production accompanied
by strict quality control. Our surveillance network in
southern China is the longest running, and the data this network
has generated have provided the most comprehensive insight
into the ecology and evolution of H5N1 virus in its natural host
(1, 2, 18?20). Despite these efforts, there remains a lack of
information in the broader region, and it is critical that similar
surveillance programs begin in other areas, including Indonesia,
Vietnam, Thailand, and India. Perhaps most importantly, information
from northern China is required, because it could answer
key questions regarding the movement of H5N1 in and out of
southern China, the hypothetical influenza epicenter (21).
Methods
Virological Surveillance, Isolation, and Characterization. Cloacal,
tracheal, and fecal samples were collected once every 7?10 days
from apparently healthy poultry in live-poultry markets in
Fujian, Guangdong, Guangxi, Guiyang, Hunan, and Yunnan
(Fig. 5). Specimens were first screened by RT-PCR for H5
subtype influenza virus. All PCR-positive swabs were shipped to
the State Key Laboratory of Emerging Infectious Diseases at
The University of Hong Kong and grown in embryonated eggs.
All isolates were identified and subtyped by using a panel of
reference antisera as previously described (22).
Antigenic Analysis. The antigenic characteristics of the H5N1
influenza viruses from different sublineages were compared by
HI assay with ferret antisera to the World Health Organization
reference H5 subtype viruses as previously described (20). The
Department of Infectious Diseases at St. Jude Children?s Research
Hospital produced the ferret antisera against DkHN
10104, IDN505, BH gooseQH1A05, and VNM120304.
Antisera to Anhui106, IDN35706, Iraq106, Whooping
swanMongolia24405, and Turkey6559606 were kindly provided
by Nancy Cox (Centers for Disease Control and Prevention,
Atlanta, GA). The HI assay started at 1:20 dilution.
To visualize similarity between the antigenic reaction patterns

 

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Re: Emergence and predominance of an H5N1 influenza

Re: Emergence and predominance of an H5N1 influenza

of different viruses, numerical analysis of HI titers was conducted
by using PRIMER version 5.2.9 (PRIMER-E, Plymouth,
United Kingdom). The data were standardized and square-roottransformed,
and the Bray?Curtis coefficient (23) was used to
construct a similarity matrix. Hierarchical agglomerative clustering
with group-average linking (24) was conducted, and a
dendrogram was produced. Nonmetric multidimensional scaling
(25) also was used to produce two- and three-dimensional
ordinations over 100 iterations. The two-dimensional configuration
with lowest overall stress was presented.
Phylogenetic Analysis and Molecular Characterization. We sequenced
the HA gene of 390 of the 1,294 (30%) H5N1 influenza
viruses isolated from poultry market surveillance in southern
China from July 2005 to June 2006 along with 16 viruses isolated
from smuggled poultry and dead wild birds in Hong Kong in
January and February 2006. In addition, 137 of those 1,294
(11%) viruses, plus the 16 Hong Kong isolates, were partially
sequenced for each of the 8 gene segments and genotyped as
previously described (1, 2, 18). Sequence assembly, editing,
alignment, and residue analysis were performed as previously
described (2). Phylogenetic analysis using MrModelTest 2.2 (26),
PAUP* 4.0 (27), and MrBayes 3.1 (28) also was carried out as
previously described (2).
Serological Analysis. Chicken sera (n
1,113) were collected from
live-poultry markets in different provinces from November 2005
to April 2006. The antibodies for H5 subtype influenza virus
were detected by HI assay, with CkHKYU2202 as the
antigen. A serum was considered positive for H5N1 virus if its
HI titer was 20. Some of those HI-positive sera were randomly
selected and further tested by neutralization assay for antibodies
against the representative isolates: CkFJ173405, CkGY
357005, and DkYN440005, which belong to the FJ-like,
GY2, and MixedVNM2 sublineages, respectively. Sera were
screened at a 1:10 series dilution against 100 TCID50 (50% tissue
culture infective dose) of those three representative viruses to
exclude negative samples, as previously described (2). Titers of
20 were regarded as positive.
We thank L. J. Zhang, J. Wong, L. Duan, and W. S. Hong for excellent
technical support. Sequence data from human cases in Indonesia were
kindly provided by the Indonesian Department of Health. This work was
supported by the Li Ka Shing Foundation and by National Institute of Allergy and Infectious Disease Contract AI95357.

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Smith et al. PNAS
November 7, 2006
vol. 103
no. 45
16941

 

[sharon sanders]

Re: PNAS - Emergence and predominance of an H5N1 influenza variant

bump this. Can't post attachments.

 

[sharon sanders]

Re: PNAS - Emergence and predominance of an H5N1 influenza variant

bump this

 

[HenryN]

Re: PNAS - Emergence and predominance of an H5N1 influenza variant

The paper includes 404 HA sequences.

Here are the 244 sequences at Los Alamos with the dominant Fujian cleavage site


DQ992831 A/chicken/Fujian/584/2006 HA (4) 1695 2006 H5N1
DQ993026 A/chicken/Guangxi/1951/2006 HA (4) 1695 2006 H5N1
DQ992745 A/chicken/Guangxi/463/2006 HA (4) 1695 2006 H5N1
DQ992770 A/chicken/Guiyang/1018/2006 HA (4) 1665 2006 H5N1
DQ992975 A/chicken/Guiyang/1211/2006 HA (4) 1059 2006 H5N1
DQ992976 A/chicken/Guiyang/1212/2006 HA (4) 1059 2006 H5N1
DQ993092 A/chicken/Guiyang/2872/2006 HA (4) 1116 2006 H5N1
DQ992763 A/chicken/Guiyang/29/2006 HA (4) 1674 2006 H5N1
DQ993094 A/chicken/Guiyang/3193/2006 HA (4) 1089 2006 H5N1
DQ993051 A/chicken/Guiyang/3194/2006 HA (4) 1059 2006 H5N1
DQ992836 A/chicken/Hong Kong/282/2006 HA (4) 1677 2006 H5N1
DQ992841 A/chicken/Hong Kong/947/2006 HA (4) 1695 2006 H5N1
ISDN184026 A/chicken/Hong Kong/947/2006 HA (4) 1706 2006 H5N1
ISDN138756 A/chicken/Malaysia/935/2006 HA (4) 1721 2006 H5N1
DQ992779 A/chicken/Shantou/1233/2006 HA (4) 1695 2006 H5N1
DQ992785 A/chicken/Shantou/3840/2006 HA (4) 1692 2006 H5N1
DQ992786 A/chicken/Shantou/3923/2006 HA (4) 1674 2006 H5N1
DQ993069 A/chicken/Yunnan/3225/2006 HA (4) 1059 2006 H5N1
DQ992843 A/common magpie/Hong Kong/2125/2006 HA (4) 1695 2006 H5N1
DQ992844 A/common magpie/Hong Kong/2256/2006 HA (4) 1695 2006 H5N1
DQ992850 A/common magpie/Hong Kong/3033/2006 HA (4) 1695 2006 H5N1
DQ992839 A/common magpie/Hong Kong/645/2006 HA (4) 1662 2006 H5N1
ISDN184030 A/Common Magpie/Hong Kong/645/2006 HA (4) 1662 2006 H5N1
DQ992838 A/crested myna/Hong Kong/540/2006 HA (4) 1671 2006 H5N1
DQ993075 A/duck/Fujian/3766/2006 HA (4) 1038 2006 H5N1
DQ993085 A/duck/Fujian/3814/2006 HA (4) 672 2006 H5N1
DQ993110 A/duck/Fujian/3834/2006 HA (4) 1068 2006 H5N1
DQ993111 A/duck/Fujian/3985/2006 HA (4) 1089 2006 H5N1
DQ993076 A/duck/Fujian/4005/2006 HA (4) 1077 2006 H5N1
DQ993084 A/duck/Fujian/5224/2006 HA (4) 693 2006 H5N1
DQ993112 A/duck/Fujian/5258/2006 HA (4) 1071 2006 H5N1
DQ993077 A/duck/Fujian/5425/2006 HA (4) 1059 2006 H5N1
DQ993113 A/duck/Fujian/5445/2006 HA (4) 1116 2006 H5N1
DQ992832 A/duck/Fujian/668/2006 HA (4) 1662 2006 H5N1
DQ992833 A/duck/Fujian/671/2006 HA (4) 1629 2006 H5N1
DQ992834 A/duck/Fujian/720/2006 HA (4) 1383 2006 H5N1
DQ993009 A/duck/Guangxi/1063/2006 HA (4) 1059 2006 H5N1
DQ992874 A/duck/Guangxi/124/2006 HA (4) 1065 2006 H5N1
DQ993030 A/duck/Guangxi/1258/2006 HA (4) 1686 2006 H5N1
DQ992875 A/duck/Guangxi/134/2006 HA (4) 1065 2006 H5N1
DQ992876 A/duck/Guangxi/143/2006 HA (4) 1059 2006 H5N1
DQ993028 A/duck/Guangxi/1436/2006 HA (4) 1683 2006 H5N1
DQ992742 A/duck/Guangxi/150/2006 HA (4) 1695 2006 H5N1
DQ993029 A/duck/Guangxi/1550/2006 HA (4) 1443 2006 H5N1
DQ993023 A/duck/Guangxi/1830/2006 HA (4) 1695 2006 H5N1
DQ993033 A/duck/Guangxi/1919/2006 HA (4) 1041 2006 H5N1
DQ993034 A/duck/Guangxi/2016/2006 HA (4) 1059 2006 H5N1
DQ993089 A/duck/Guangxi/2042/2006 HA (4) 1116 2006 H5N1
DQ992878 A/duck/Guangxi/264/2006 HA (4) 1059 2006 H5N1
DQ992744 A/duck/Guangxi/288/2006 HA (4) 1695 2006 H5N1
DQ992879 A/duck/Guangxi/309/2006 HA (4) 1056 2006 H5N1
DQ993013 A/duck/Guangxi/340/2006 HA (4) 1059 2006 H5N1
DQ993015 A/duck/Guangxi/508/2006 HA (4) 1059 2006 H5N1
DQ992748 A/duck/Guangxi/619/2006 HA (4) 1647 2006 H5N1
DQ992749 A/duck/Guangxi/744/2006 HA (4) 1647 2006 H5N1
DQ992750 A/duck/Guangxi/804/2006 HA (4) 1647 2006 H5N1
DQ993019 A/duck/Guiyang/1081/2006 HA (4) 1059 2006 H5N1
DQ992773 A/duck/Guiyang/1260/2006 HA (4) 1665 2006 H5N1
DQ992977 A/duck/Guiyang/1261/2006 HA (4) 1059 2006 H5N1
DQ992978 A/duck/Guiyang/1262/2006 HA (4) 1059 2006 H5N1
DQ992980 A/duck/Guiyang/1417/2006 HA (4) 1059 2006 H5N1
DQ992775 A/duck/Guiyang/1418/2006 HA (4) 1695 2006 H5N1
DQ992981 A/duck/Guiyang/1441/2006 HA (4) 1059 2006 H5N1
DQ992982 A/duck/Guiyang/1450/2006 HA (4) 1059 2006 H5N1
DQ992984 A/duck/Guiyang/1557/2006 HA (4) 1059 2006 H5N1
DQ992985 A/duck/Guiyang/1558/2006 HA (4) 1059 2006 H5N1
DQ992986 A/duck/Guiyang/1596/2006 HA (4) 1059 2006 H5N1
DQ992992 A/duck/Guiyang/1705/2006 HA (4) 1059 2006 H5N1
DQ992993 A/duck/Guiyang/1722/2006 HA (4) 1059 2006 H5N1
DQ992994 A/duck/Guiyang/1739/2006 HA (4) 1059 2006 H5N1
DQ993040 A/duck/Guiyang/2044/2006 HA (4) 1071 2006 H5N1
DQ993042 A/duck/Guiyang/2159/2006 HA (4) 1041 2006 H5N1
DQ993090 A/duck/Guiyang/2199/2006 HA (4) 1083 2006 H5N1
DQ993078 A/duck/Guiyang/2453/2006 HA (4) 840 2006 H5N1
DQ993044 A/duck/Guiyang/2489/2006 HA (4) 1059 2006 H5N1
DQ993046 A/duck/Guiyang/2637/2006 HA (4) 1071 2006 H5N1
DQ993091 A/duck/Guiyang/2647/2006 HA (4) 1101 2006 H5N1
DQ993047 A/duck/Guiyang/2757/2006 HA (4) 1071 2006 H5N1
DQ992914 A/duck/Guiyang/285/2006 HA (4) 702 2006 H5N1
DQ993096 A/duck/Guiyang/3210/2006 HA (4) 1083 2006 H5N1
DQ993079 A/duck/Guiyang/3215/2006 HA (4) 720 2006 H5N1
DQ993053 A/duck/Guiyang/3353/2006 HA (4) 1059 2006 H5N1
DQ993055 A/duck/Guiyang/3515/2006 HA (4) 1059 2006 H5N1
DQ992767 A/duck/Guiyang/497/2006 HA (4) 1695 2006 H5N1
DQ992998 A/duck/Hunan/1152/2006 HA (4) 1059 2006 H5N1
DQ992999 A/duck/Hunan/1159/2006 HA (4) 1059 2006 H5N1
DQ993000 A/duck/Hunan/1172/2006 HA (4) 1059 2006 H5N1
DQ993001 A/duck/Hunan/1186/2006 HA (4) 1059 2006 H5N1
DQ993002 A/duck/Hunan/1204/2006 HA (4) 1059 2006 H5N1
DQ993003 A/duck/Hunan/1213/2006 HA (4) 1059 2006 H5N1
DQ993004 A/duck/Hunan/1231/2006 HA (4) 1059 2006 H5N1
DQ993005 A/duck/Hunan/1247/2006 HA (4) 1059 2006 H5N1
DQ993006 A/duck/Hunan/1260/2006 HA (4) 1059 2006 H5N1
DQ993007 A/duck/Hunan/1271/2006 HA (4) 1056 2006 H5N1
DQ993008 A/duck/Hunan/1286/2006 HA (4) 1059 2006 H5N1
DQ992933 A/duck/Hunan/301/2006 HA (4) 1059 2006 H5N1
DQ992934 A/duck/Hunan/307/2006 HA (4) 1059 2006 H5N1
DQ992935 A/duck/Hunan/319/2006 HA (4) 1059 2006 H5N1
DQ992790 A/duck/Hunan/324/2006 HA (4) 1647 2006 H5N1
DQ992936 A/duck/Hunan/336/2006 HA (4) 1059 2006 H5N1
DQ992791 A/duck/Hunan/344/2006 HA (4) 1647 2006 H5N1
DQ992937 A/duck/Hunan/350/2006 HA (4) 1020 2006 H5N1
DQ992938 A/duck/Hunan/364/2006 HA (4) 1059 2006 H5N1
DQ992939 A/duck/Hunan/382/2006 HA (4) 948 2006 H5N1
DQ992940 A/duck/Hunan/391/2006 HA (4) 696 2006 H5N1
DQ992792 A/duck/Hunan/856/2006 HA (4) 1692 2006 H5N1
DQ993012 A/duck/Hunan/867/2006 HA (4) 1059 2006 H5N1
DQ992793 A/duck/Hunan/988/2006 HA (4) 1611 2006 H5N1
DQ993010 A/duck/Hunan/995/2006 HA (4) 1059 2006 H5N1
DQ845348 A/duck/Laos/3295/2006 HA (4) 1719 2006 H5N1
ISDN138780 A/duck/Laos/3295/2006 HA (4) 1719 2006 H5N1
DQ993098 A/duck/Yunnan/2075/2006 HA (4) 1071 2006 H5N1
DQ993099 A/duck/Yunnan/2099/2006 HA (4) 1089 2006 H5N1
DQ993115 A/duck/Yunnan/2107/2006 HA (4) 1071 2006 H5N1
DQ993116 A/duck/Yunnan/2113/2006 HA (4) 1056 2006 H5N1
DQ993100 A/duck/Yunnan/2267/2006 HA (4) 924 2006 H5N1
DQ993101 A/duck/Yunnan/2268/2006 HA (4) 888 2006 H5N1
DQ993102 A/duck/Yunnan/2272/2006 HA (4) 930 2006 H5N1
DQ993062 A/duck/Yunnan/2273/2006 HA (4) 1059 2006 H5N1
DQ993065 A/duck/Yunnan/2535/2006 HA (4) 1074 2006 H5N1
DQ993081 A/duck/Yunnan/2579/2006 HA (4) 699 2006 H5N1
DQ993109 A/duck/Yunnan/3278/2006 HA (4) 1083 2006 H5N1
DQ993070 A/duck/Yunnan/3299/2006 HA (4) 1065 2006 H5N1
DQ993071 A/duck/Yunnan/3332/2006 HA (4) 1071 2006 H5N1
DQ993027 A/goose/Guangxi/1458/2006 HA (4) 1695 2006 H5N1
DQ993025 A/goose/Guangxi/1633/2006 HA (4) 1695 2006 H5N1
DQ993032 A/goose/Guangxi/1786/2006 HA (4) 1038 2006 H5N1
DQ993024 A/goose/Guangxi/1898/2006 HA (4) 1695 2006 H5N1
DQ992877 A/goose/Guangxi/222/2006 HA (4) 1062 2006 H5N1
DQ992743 A/goose/Guangxi/224/2006 HA (4) 1632 2006 H5N1
DQ992873 A/goose/Guangxi/30/2006 HA (4) 1065 2006 H5N1
DQ992740 A/goose/Guangxi/52/2006 HA (4) 1665 2006 H5N1
DQ993117 A/goose/Guangxi/532/2006 HA (4) 1695 2006 H5N1
DQ993021 A/goose/Guiyang/1147/2006 HA (4) 1059 2006 H5N1
DQ992774 A/goose/Guiyang/1304/2006 HA (4) 1695 2006 H5N1
DQ992983 A/goose/Guiyang/1461/2006 HA (4) 1059 2006 H5N1
DQ992987 A/goose/Guiyang/1609/2006 HA (4) 1059 2006 H5N1
DQ992988 A/goose/Guiyang/1636/2006 HA (4) 1059 2006 H5N1
DQ992989 A/goose/Guiyang/1639/2006 HA (4) 1059 2006 H5N1
DQ992990 A/goose/Guiyang/1647/2006 HA (4) 1059 2006 H5N1
DQ992995 A/goose/Guiyang/1785/2006 HA (4) 1059 2006 H5N1
DQ992996 A/goose/Guiyang/1794/2006 HA (4) 1059 2006 H5N1
DQ992997 A/goose/Guiyang/1796/2006 HA (4) 1059 2006 H5N1
DQ993041 A/goose/Guiyang/2071/2006 HA (4) 1071 2006 H5N1
DQ993043 A/goose/Guiyang/2231/2006 HA (4) 1059 2006 H5N1
DQ993086 A/goose/Guiyang/2368/2006 HA (4) 678 2006 H5N1
DQ993045 A/goose/Guiyang/2502/2006 HA (4) 1041 2006 H5N1
DQ993048 A/goose/Guiyang/2839/2006 HA (4) 1068 2006 H5N1
DQ993050 A/goose/Guiyang/2988/2006 HA (4) 1071 2006 H5N1
DQ993052 A/goose/Guiyang/3283/2006 HA (4) 1059 2006 H5N1
DQ993054 A/goose/Guiyang/3435/2006 HA (4) 1059 2006 H5N1
DQ992917 A/goose/Guiyang/380/2006 HA (4) 1062 2006 H5N1
DQ992768 A/goose/Guiyang/538/2006 HA (4) 1143 2006 H5N1
DQ993018 A/goose/Guiyang/765/2006 HA (4) 1059 2006 H5N1
DQ992783 A/goose/Shantou/3265/2006 HA (4) 1686 2006 H5N1
DQ992924 A/goose/Shantou/3285/2006 HA (4) 777 2006 H5N1
DQ992784 A/goose/Shantou/3295/2006 HA (4) 1695 2006 H5N1
DQ992815 A/goose/Yunnan/1143/2006 HA (4) 1674 2006 H5N1
DQ992816 A/goose/Yunnan/1144/2006 HA (4) 1695 2006 H5N1
DQ992966 A/goose/Yunnan/1155/2006 HA (4) 1059 2006 H5N1
DQ992818 A/goose/Yunnan/1396/2006 HA (4) 1695 2006 H5N1
DQ993059 A/goose/Yunnan/1770/2006 HA (4) 1056 2006 H5N1
DQ993097 A/goose/Yunnan/1779/2006 HA (4) 1071 2006 H5N1
DQ993060 A/goose/Yunnan/1798/2006 HA (4) 1074 2006 H5N1
DQ993061 A/goose/Yunnan/2140/2006 HA (4) 1020 2006 H5N1
DQ993105 A/goose/Yunnan/2190/2006 HA (4) 1065 2006 H5N1
DQ993103 A/goose/Yunnan/2337/2006 HA (4) 858 2006 H5N1
DQ993104 A/goose/Yunnan/2338/2006 HA (4) 930 2006 H5N1
DQ993063 A/goose/Yunnan/2398/2006 HA (4) 1068 2006 H5N1
DQ993064 A/goose/Yunnan/2485/2006 HA (4) 1077 2006 H5N1
DQ993106 A/goose/Yunnan/2501/2006 HA (4) 1089 2006 H5N1
DQ993107 A/goose/Yunnan/2512/2006 HA (4) 1101 2006 H5N1
DQ993066 A/goose/Yunnan/2739/2006 HA (4) 1056 2006 H5N1
DQ993067 A/goose/Yunnan/2761/2006 HA (4) 1047 2006 H5N1
DQ993072 A/goose/Yunnan/3340/2006 HA (4) 1077 2006 H5N1
DQ993082 A/goose/Yunnan/3359/2006 HA (4) 732 2006 H5N1
DQ993073 A/goose/Yunnan/3479/2006 HA (4) 1059 2006 H5N1
DQ992848 A/house crow/Hong Kong/2648/2006 HA (4) 1695 2006 H5N1
DQ992849 A/house crow/Hong Kong/2858/2006 HA (4) 1695 2006 H5N1
ISDN184028 A/Japanese White Eye/Hong Kong/1038/2006 HA (4) 1693 2006 H5N1
DQ992842 A/Japanese white-eye/Hong Kong/1038/2006 HA (4) 1692 2006 H5N1
DQ992847 A/large-billed crow/Hong Kong/2512/2006 HA (4) 1695 2006 H5N1
DQ992840 A/little egret/Hong Kong/718/2006 HA (4) 1692 2006 H5N1
DQ992845 A/munia/Hong Kong/2454/2006 HA (4) 1692 2006 H5N1
ISDN184024 A/Munia/Hong Kong/2454/2006 HA (4) 1693 2006 H5N1
DQ992837 A/robin/Hong Kong/366/2006 HA (4) 1500 2006 H5N1
DQ992835 A/robin/Hong Kong/75/2006 HA (4) 1692 2006 H5N1
DQ992846 A/white-backed munia/Hong Kong/2469/2006 HA (4) 1695 2006 H5N1
DQ643809 A/Zhejiang/16/2006 HA (4) 1281 2006 H5N1
DQ371928 A/Anhui/1/2005 HA (4) 1704 2005 H5N1
DQ371929 A/Anhui/2/2005 HA (4) 1704 2005 H5N1
DQ992828 A/chicken/Fujian/11933/2005 HA (4) 1695 2005 H5N1
DQ992830 A/chicken/Fujian/12239/2005 HA (4) 1680 2005 H5N1
DQ992734 A/chicken/Guangxi/4989/2005 HA (4) 1677 2005 H5N1
DQ992759 A/chicken/Guiyang/3721/2005 HA (4) 1695 2005 H5N1
DQ992903 A/chicken/Guiyang/3923/2005 HA (4) 1059 2005 H5N1
DQ992762 A/chicken/Guiyang/4059/2005 HA (4) 1677 2005 H5N1
DQ992971 A/duck/Fujian/10160/2005 HA (4) 1044 2005 H5N1
DQ095629 A/Duck/Fujian/1734/05 HA (4) 1703 2005 H5N1
DQ992859 A/duck/Guangxi/4409/2005 HA (4) 1065 2005 H5N1
DQ992730 A/duck/Guangxi/4428/2005 HA (4) 1695 2005 H5N1
DQ992861 A/duck/Guangxi/4561/2005 HA (4) 1065 2005 H5N1
DQ992862 A/duck/Guangxi/4686/2005 HA (4) 1062 2005 H5N1
DQ992863 A/duck/Guangxi/4823/2005 HA (4) 1065 2005 H5N1
DQ992733 A/duck/Guangxi/4830/2005 HA (4) 1695 2005 H5N1
DQ992864 A/duck/Guangxi/4836/2005 HA (4) 1065 2005 H5N1
DQ992868 A/duck/Guangxi/4923/2005 HA (4) 1065 2005 H5N1
DQ992735 A/duck/Guangxi/5075/2005 HA (4) 1647 2005 H5N1
DQ992736 A/duck/Guangxi/5165/2005 HA (4) 1695 2005 H5N1
DQ992869 A/duck/Guangxi/5195/2005 HA (4) 1065 2005 H5N1
DQ992737 A/duck/Guangxi/5270/2005 HA (4) 1695 2005 H5N1
DQ992871 A/duck/Guangxi/5430/2005 HA (4) 1059 2005 H5N1
DQ992872 A/duck/Guangxi/5449/2005 HA (4) 1062 2005 H5N1
DQ992739 A/duck/Guangxi/5457/2005 HA (4) 1695 2005 H5N1
DQ992882 A/duck/Guiyang/3026/2005 HA (4) 1026 2005 H5N1
DQ992897 A/duck/Guiyang/3664/2005 HA (4) 1059 2005 H5N1
DQ992760 A/duck/Guiyang/3834/2005 HA (4) 1647 2005 H5N1
DQ992761 A/duck/Guiyang/3996/2005 HA (4) 1689 2005 H5N1
DQ992926 A/duck/Hunan/5102/2005 HA (4) 1059 2005 H5N1
DQ992787 A/duck/Hunan/5106/2005 HA (4) 1695 2005 H5N1
DQ992927 A/duck/Hunan/5118/2005 HA (4) 792 2005 H5N1
DQ992928 A/duck/Hunan/5128/2005 HA (4) 1059 2005 H5N1
DQ992929 A/duck/Hunan/5146/2005 HA (4) 1059 2005 H5N1
DQ992788 A/duck/Hunan/5152/2005 HA (4) 1695 2005 H5N1
DQ992930 A/duck/Hunan/5170/2005 HA (4) 1059 2005 H5N1
DQ992931 A/duck/Hunan/5180/2005 HA (4) 819 2005 H5N1
DQ992932 A/duck/Hunan/5191/2005 HA (4) 1059 2005 H5N1
DQ992789 A/duck/Hunan/5472/2005 HA (4) 1068 2005 H5N1
DQ992776 A/duck/Shantou/13323/2005 HA (4) 1695 2005 H5N1
DQ992856 A/goose/Guangxi/4274/2005 HA (4) 1065 2005 H5N1
DQ992729 A/goose/Guangxi/4289/2005 HA (4) 1680 2005 H5N1
DQ992731 A/goose/Guangxi/4513/2005 HA (4) 1695 2005 H5N1
DQ992865 A/goose/Guangxi/4878/2005 HA (4) 1062 2005 H5N1
DQ992866 A/goose/Guangxi/4881/2005 HA (4) 1062 2005 H5N1
DQ992867 A/goose/Guangxi/4904/2005 HA (4) 1065 2005 H5N1
DQ992900 A/goose/Guiyang/3765/2005 HA (4) 1059 2005 H5N1
DQ992906 A/goose/Guiyang/4006/2005 HA (4) 1017 2005 H5N1
DQ992907 A/goose/Guiyang/4021/2005 HA (4) 1059 2005 H5N1
DQ992908 A/goose/Guiyang/4030/2005 HA (4) 1059 2005 H5N1
DQ992921 A/goose/Shantou/18428/2005 HA (4) 1062 2005 H5N1
DQ992777 A/goose/Shantou/18442/2005 HA (4) 1647 2005 H5N1
DQ992922 A/goose/Shantou/18446/2005 HA (4) 1062 2005 H5N1
DQ992809 A/goose/Yunnan/6169/2005 HA (4) 1623 2005 H5N1
DQ371930 A/Guangxi/1/2005 HA (4) 1704 2005 H5N1