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PostPosted: Wed Dec 24, 2014 5:22 am 
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Recent Veterinary Research paper describes receptor binding properties for H5 clade 2.3.4.6 (same Fujian sub-clade as H5N8 and H5N2 in Europe, Japan, and North America.

http://www.veterinaryresearch.org/content/45/1/127

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PostPosted: Wed Dec 24, 2014 5:31 am 
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Novel H5 clade 2.3.4.6 viruses with both α-2,3 and α-2,6 receptor binding properties may pose a pandemic threat
Qunhui Li1, Xuan Wang1, Min Gu12, Jie Zhu1, Xiaoli Hao1, Zhao Gao1, Zhongtao Sun1, Jiao Hu12, Shunlin Hu12, Xiaoquan Wang12, Xiaowen Liu12 and Xiufan Liu12*

* Corresponding author: Xiufan Liu xfliu@yzu.edu.cn

Author Affiliations
1 Animal Infectious Disease Laboratory, College of Veterinary Medicine, Yangzhou University, Yangzhou 225009, Jiangsu, China

2 Jiangsu Co-innovation Center for Prevention and Control of Important Animal Infectious Diseases and Zoonoses, Yangzhou 225009, Jiangsu, China


Received: 15 September 2014
Accepted: 1 December 2014
Published: 17 December 2014

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PostPosted: Wed Dec 24, 2014 5:32 am 
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Abstract
The emerging H5 clade 2.3.4.6 viruses of different NA subtypes have been detected in different domestic poultry in China. We evaluated the receptor binding property and transmissibility of four novel H5 clade 2.3.4.6 subtype highly pathogenic avian influenza viruses. The results show that these viruses bound to both avian-type (α-2,3) and human-type (α-2,6) receptors. Furthermore, we found that one of these viruses, GS/EC/1112/11, not only replicated but also transmitted efficiently in guinea pigs. Therefore, such novel H5 subtype viruses have the potential of a pandemic threat.

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PostPosted: Wed Dec 24, 2014 11:02 am 
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Introduction, methods, and results
H5N1 subtype highly pathogenic avian influenza virus (HPAIV) was first isolated in sick geese in China in 1996,and has continued to evolve into over 10 distinct phylogenetic clades including different subclades based on the hemagglutinin (HA) gene [1]. Since 2010, H5 HPAIV subtypes which belong to the recommended novel clade 2.3.4.6 [2] with various neuraminidase (NA) subtypes (H5N1, H5N2, H5N6 and H5N8) have been detected in different domestic poultry in China [2]-[7]. Furthermore, the H5N8 virus-caused outbreaks have also been reported in wild birds and poultry in South Korea and Japan in January and April, 2014 respectively [8],[9]. Here, we tested the receptor binding property of four novel clade 2.3.4.6 viruses, and guinea pigs were used as a mammalian model to examine the replication and transmission of these viruses. All animal experiments were approved by the Jiangsu Administrative Committee for Laboratory Animals (permission number SYXK-SU-2007-0005) and complied with the guidelines of the Jiangsu laboratory animal welfare and ethics of Jiangsu Administrative Committee of Laboratory Animals.

During surveillance of poultry for avian influenza viruses in live poultry markets in eastern China in 2013, one H5N8 avian influenza virus, A/duck/Shandong/Q1/2013 (DkQ1), was isolated from domestic ducks. The GenBank accession numbers for the DkQ1 segments are KM504098 to KM504105. Sequence analysis showed that all 8 genes of DkQ1 are closely related to those H5N8 viruses which have been reported in eastern China [4],[5]. Furthermore, the HA gene of DkQ1 has high nucleotide identity with the H5N8 viruses circulating in South Korea and Japan in 2014 [8],[9]. And all these H5N8 viruses belong to the recommended novel clade 2.3.4.6 [2]. In addition, one H5N8 virus A/duck/Jiangsu/k1203/2010 (Dkk1203) [4] and two H5N2 viruses A/duck/Eastern China/1111/2011 (DK/EC/1111/11) and A/goose/Eastern China/1112/2011 (GS/EC/1112/11) [3], which have been reported to circulate in eastern China, also possess HA genes belonging to the novel clade 2.3.4.6. Here, we investigated the receptor binding property and transmissibility of these four H5 (HPAIV) clade 2.3.4.6. All experiments with viruses were performed in a Biosafety Level 3 laboratory.

It is generally accepted that haemagglutinin-receptor-binding preference to α-2,6-linked sialylated glycans is the initial key step for a novel influenza-virus-causing pandemic [10]. First, we examined the receptor-binding specificity of these reassortant viruses by hemagglutination assays using goose red blood cells that were treated with a α-2,3-specific sialidase as previously described [11]. The A(H1N1)pdm2009 virus A/California/04/2009 (CA/04) and poultry H5N1 isolate A/mallard/Huadong/S/2005 (HD/05) [12] were used as controls. Theoretically, the sialidase digestion should abolish hemagglutination by α-2,3-specific viruses, whereas viruses that can bind to α-2,6-receptors should maintain hemagglutination activity with the treated red blood cells. The sialidase treatment did not affect the hemagglutination titer of CA/04, as shown in Table 1. Compared to untreated GRBC, these reassortant viruses still show some lower HA activity with α-2,3-sialidase-treated GRBC, which had only α-2,6-receptors (Table 1).

Table 1. Hemagglutination titers of viruses from humans and animalsa
To characterize the receptor-binding properties of these viruses further, we performed solid-phase binding assays with different glycans as previously described [13]. Briefly, the synthetic sialylglycopolymers Neu5Aca2-3Galb1-4GlcNAcb (3’SLN)-PAA-biotin and Neu5Aca2-3Galb1-4GlcNAcb (6’SLN)-PAA-biotin (GlycoTech) were serially diluted in PBS and added to the wells of 96-well streptavidin coated microtiter plates (Pierce). The plates were blocked with PBS containing 2% skim milk powder, and 128 HA units of live virus was added per well. Chicken antiserum against the virus was diluted in PBS and added to each well. Bound antibody was detected by sequential addition of HRP-conjugated rabbit anti-chicken IgG antibody and tetramethylbenzidine substrate solution. The reaction was stopped with 1 M H2SO4, and the absorbance was read at 450 nm. Each sample was measured in triplicate. Our results show these reassortant viruses were bound to both avian-type (α-2,3) and human-type (α-2,6) receptors, whereas HD/05 and CA/04 viruses were preferentially bound to α-2,3 and α-2,6 receptors respectively, as expected (Figure 1). These results indicate that the HA of these reassortant viruses binds to α-2,3 receptors as well as to α-2,6 receptors.

thumbnailFigure 1. Solid-phase receptor-binding assay of the H5 (HPAIV) clades 2.3.4.6. Solid-phase receptor-binding assay of human isolate CA/04 (A), poultry isolate HD/05 (B), DK/EC/1111/11 virus (C), GS/EC/1112/11 virus (D), Dkk1203 virus (E) and DkQ1 virus (F). Direct binding of viruses to sialylglycopolymers containing either 3’SLN-PAA or 6’SLN-PAA was measured. The data shown are representative of three independent binding experiments.
To investigate the replication of these reassortant viruses, groups of four animals were anesthetized with pentobarbital natricum (40–50 mg/Kg) and inoculated intranasally with 106EID50 of test virus in a 300 μL volume (150 μL per nostril). Two animals from each group were euthanized with CO2 on day 3 post inoculation (pi) and nasal washes, tracheas, lungs, kidneys, spleens, and brains were collected for virus titration in eggs. The remaining two animals were observed for two weeks for signs of disease and death. The A(H1N1)pdm2009 virus A/California/04/2009 (CA/04) and poultry H5N1 isolate A/mallard/Huadong/S/2005 (HD/05) were used as controls. As shown in Table 2, all reassortant viruses were detected in the nasal washes, tracheas and lungs of both inoculated animals, but only could be detected at lower titers in the trachea and lungs of infected guinea pigs. Virus was not detected in the brains, kidneys or spleens of any of the inoculated animals. We also infected two animals for each virus and observed them for two weeks for signs of pathogenicity. After two weeks pi, all of the animals seroconverted (Table 2). None of the animals showed disease signs during the observation period. These results indicate that replication of these reassortant viruses in guinea pigs is restricted to the respiratory system.

Table 2. Virus replication and seroconversion in guinea pigs
For the contact transmission studies, groups of three animals were inoculated intranasally with 106EID50 of test virus and housed in a cage placed inside an isolator. Three naïve animals were introduced into the same cage 24 h later. Nasal washes were collected at 2 day intervals, beginning on day 2 pi (1 day post contact) and titrated in eggs. Sera were collected from guinea pigs at 14 days post inoculation (dpi) for hemagglutinin inhibition (HI) antibody detection [14]. Evidence of transmission was based on the detection of virus in the nasal wash and on seroconversion at the end of the two-week observation period. The A/California/04/2009 (CA/04) virus was used as controls. As shown in Figure 2, reassortant virus was detected in the nasal washes of all three inoculated guinea pigs between days 2–6 pi, but not in any of the contact guinea pigs. In the GS/EC/1112/11-inoculated groups (Figure 2B), virus was detected in the nasal washes of all three inoculated guinea pigs between days 2–6 pi, respectively and was also detected in the nasal washes of all three contact animals between days 4–8 pi. Seroconversion occurred in all inoculated groups (Table 2). In the contact animal groups, seroconversion was only observed among animals placed with the GS/EC/1112/11-inoculated animals. These results indicate that the transmissibility of the reassortant viruses in guinea pigs varies among viral strains, and of the four test viruses, only GS/EC/1112/11 transmit efficiently in this mammalian host.

thumbnailFigure 2. Transmisson of the H5 (HPAIV) clades 2.3.4.6 in guinea pigs. (A) DK/EC/1111/11 virus, (B) GS/EC/1112/11 virus, (C) Dkk1203 virus, (D) DkQ1 virus and (E) CA/04 virus. Each color bar represents the virus titer from an individual animal. The dashed blue lines in these panels indicate the lower limit of detection.

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PostPosted: Wed Dec 24, 2014 11:17 am 
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Discussion
Historically, changes in the receptor binding protein of influenza virus, HA, have been implicated in the initiation of a pandemic. It has been established for the H1N1 (1918), H2N2 (1957) and H3N2 (1968) pandemic viruses that a change in HA protein from a preference for α-2,3-linked sialic acids (avian receptor) to a preference for α-2,6-linked sialic acids (human receptor) is a prerequisite for efficient transmission of avian viruses to humans [10]. H5 HPAIV pose a serious pandemic threat due to their virulence and high mortality in humans, and their increasingly expanding host reservoir and significant ongoing evolution could enhance their human-to-human transmissibility. Recently, novel clade 2.3.4.6 H5 HPAIV with various NA subtypes (H5N1, H5N2, H5N6, and H5N8) were reported in Eastern China and South Korea [2]-[7],[9],[15]. Here, we evaluated their receptor specificity and transmission in guinea pigs. The results show that the viruses bound to both avian-type (α-2,3) and human-type (α-2,6) receptors. In humans, the α-2,6 receptor is expressed mainly in the upper airway, while the α-2,3 receptor is expressed in alveoli and the terminal bronchiole [16]. A virus with good affinity to both α-2,3 and α-2,6 receptors may especially be harmful, as it could infect efficiently via its binding to α-2,6 receptors in the upper airway and simultaneously cause severe infection in the lung via its binding to α-2,3 receptors. And this hypothesis is supported by the fact that one of the two well-characterized HA genes from the H1N1 1918 pandemic virus binds efficiently to both α-2,3 and α-2,6 receptors [17]. In addition, previous studies showed that the human-infecting novel H7N9 and the latest reassortant H10N8 avian influenza viruses yet have substantial affinity to both avian-type (α-2,3) and human-type (α-2,6) receptors [18],[19]. Sequence analysis showed that novel H5 (HPAIV) clade 2.3.4.6 simultaneously carry a T160A mutation which results in the lack of an oligosaccharide side chain at 158–160 of HA, and it is critical for the H5 subtype influenza viruses tested to bind to human-like receptors and to transmit among a mammalian host [20],[21]. Whether this T160A variation affects the receptor-binding property deserves further investigation. Previous studies showed that some H5 subtype influenza viruses can transmit efficiently in guinea pigs [21]. In this study, we also found that one of these viruses, GS/EC/1112/11, not only replicated but also transmitted efficiently in guinea pigs. These findings emphasize that continued circulation of these viruses may pose health threats for humans. Therefore, we need to intensify our effort to detect such viruses as early as possible.

Abbreviations
HPAIV: Highly pathogenic avian influenza virus

HA: Hemagglutinin

NA: Neuraminidase

DkQ1: A/duck/Shandong/Q1/2013

Dkk1203: A/duck/Jiangsu/k1203/2010

DK/EC/1111/11: A/duck/Eastern China/1111/2011

GS/EC/1112/11: A/goose/Eastern China/1112/2011

CA/04: A/California/04/2009

HD/05: A/mallard/Huadong/S/2005

pi: Post inoculation

dpi: Days post inoculation

HI: Hemagglutinin inhibition

Competing interests
The authors declare that they have no competing interests.

Authors’ contributions
QL designed the study, drafted the manuscript and participated in all tests. XW verified design, participated in all tests. JZ participated in collecting samples. XH, ZS and ZG participated in testing samples. MG, JH, SH, XQW, XWL and XFL conceived the study, contributed to the analysis of the results and preparation of revised manuscript versions. All authors read and approved the final manuscript.

Acknowledgments
This work was supported by the Earmarked Fund for Modern Agro-Industry Technology Research System (grant no. nycytx-41-G07), the Jiangsu Provincial Natural Science Foundation of China (grant no. BK20130442), the Priority Academic Program Development of Jiangsu Higher Education Institutions (PAPD), the National Natural Science Foundation of China (grant no. 31101827), and the National High-Tech R&D Program of China (863 Program) (grant no. 2011AA10A200).

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PostPosted: Wed Dec 24, 2014 11:18 am 
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PostPosted: Thu Jan 15, 2015 12:51 pm 
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Real threat of avian flu spreading to humans

By Wong Ruey-hong 翁瑞宏
Avian influenza continues to spread. Following an outbreak of the H5N2 strain in Pingtung County, a new strain of H5N2 and an H5N8 strain that had never before been seen domestically have been found in central and southern Taiwan.
The Council of Agriculture’s Bureau of Animal and Plant Health Inspection and Quarantine said that the H5N2 subtype cannot be spread from animals to humans and that the virus does not have any negative effects on the human body.
The question is: Is this really true?
In 2005, in Ibaraki Prefecture in Japan, 5.78 million chickens at 41 farms were infected with the low-pathogenic H5N2 virus.
Although no one who had been in contact with the chickens showed any symptoms, the result of a serological study found that workers at poultry farms had higher levels of H5N2 antibodies than healthy people in the general population. It is important to note that seasonal flu vaccine inoculations or other anti-flu treatments might have an influence on neutralizing antibody-positive status. However, this study has been seen as the first example of people being infected with H5N2.
Last year, a group of researchers in Niger published a report saying that they had found H5N2 antibodies in eight people who had been in contact with animals. In Taiwan, avian infections with low-pathogenic H5N2 were reported in 2003 and 2004, as well as between 2008 and 2011, while infections with the highly pathogenic H5N2 subtype occurred for the first time in 2012.
Following the 2012 epidemic, six people who had been in contact with animals were discovered by staff from the Centers for Disease Control to have antibodies, which means that they could have been infected with H5N2.
Furthermore, a research team in South Korea discovered the H5N2 virus in pigs, a result that shows that interspecies infections from poultry to mammals has already occurred. This is more evidence that the H5N2 virus might be capable of genetic recombination and could become highly pathogenic to humans.
Last year, South Korea set a new record when it culled 15 million domestic birds due to another highly pathogenic strain, H5N8. South Korean researchers issued an urgent public health alert because they found that this new virus contained the highly pathogenic antigen H5, that the virus could bind to human-like receptors and that it could reproduce in the respiratory tracts of mammals.
Based on this academic evidence, it can be said that although researchers still cannot draw the conclusion that the H5N2 and H5N8 subtypes infect people, there is the possibility that they can.
In particular, flu epidemics are most common during the winter, and once a human influenza virus comes in contact with the avian flu virus, it is possible that genetic recombination could occur. This is why the government must not take lightly the occurrence of the H5N2 and H5N8 viruses, and it is imperative that they maintain strict control of epidemics and quickly cull any poultry that has been infected.
In addition, if anyone who has been in contact with poultry or pigs develops a fever, a cough or muscle pain, they must wear a face mask and immediately contact a doctor, remembering to describe the circumstances under which they were in contact with the animal.
At the same time, the public should be sure that chicken and eggs are thoroughly cooked before eating.

Wong Ruey-hong is a professor in the School of Public Health at Chung Shan Medical University.
Translated by Perry Svensson

http://www.taipeitimes.com/News/editori ... 2003609341

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