13340 ﾒ. 6ｾ409. ﾒ0806ｦﾐ ﾞ ﾒ ﾇ6ｾ4, VAN REETH K ｦ ｧ9000ｹ0: ( ) 02080ｸ70ｹ ｦ00ｹ ｧ80ｹ fi07080

Transcription

1 13096ﾑ806ｦﾐ ﾌ04ｦﾐ ﾒ ﾌ 6ﾒ2 ﾇ6ｾ4 E HNIKH6ｾ4 KTHNｦﾐ ﾞ6ﾒ206ｦﾐ ﾒ ﾇ6ｾ4 ETAIP6ﾑ8IA6ｾ4 2005, 56(4): : ﾑ fi Special Article H 09036ｧ9apple09 020ｷ307 apple : 08 03fi03080ｹ ｸ ｧ903080ｸ ｧ907 0ｸ fi0406 0ｸ709006ｧ9 ﾒ0ｷ307. 6ｾ4apple. ﾒ0ｸ ｹ0, Van Reeth K. ﾞvian influenza: the role of the pig and public health 06mplications Kyriakis C. S., Van Reeth K. 096ﾑ806ｦﾐ ﾇ03 ﾇ. ﾇ apple06ｨB0ｷ308026ｧ ｹ ｧ9apple090ｹ0 020ｷ307 apple , ｧ9 0ｹ ｸ 閨04080ｸ70ｹ0 0ｸ ｧ80ｹ fi07080ｸ fi ｹ0 ﾇ5031, ｧ8 ﾞ046ｧ ｸ702 6ｧ fi0706, ｸ7036ｧ90ｷ30ｹ ｦ ｧ9apple090ｹ0 020ｷ307 apple apple080ｸ70ｹ0 閨04080ｸ 閨 ｸ fi ｧ8 00apple ｧ fi apple fi ｧ8030ｷ304090ｹ ｹ0 07 陦 0ｹ0 apple ｧ9 0ｹ ｧ9apple090ｹ0: apple ｸ ｸ ｾ ｸ ｸ ｷ3applefi apple apple ｧ9000ｹ0, ｸ70ｹ ｦﾐ04apple ｧ ｧ9apple ｸ , apple080ｸ7 0402fi020ｷ30400 apple fi ｸ70ｹ0 applefi ｦ apple080ｸ70ｹ0 04 ｯfi fi ﾌ06 00applefi ｹ0 6ｦ008 apple ｧ9000ｹ0 0ｸ7apple6ｧ fi 閨0ｹ0, ｸ apple00036ｧ9apple080ｸ ｦ ｦ ﾌ ｧ ｹ ｧ9apple090ｹ ｸ7 026ｦ0apple080ｸ7 ﾞ 076ｧ802080ｸ ｸ70ｹ0 ﾌ ｸ8086ｦ00ｹ0, ｷ ｧ ｸ70ｹ0 006ｧ 閨 fi02030ｷ30708 RNA. Ta RNA 02080ｸ ｦ0 020ｷ apple ｦ apple030ｷ302000ｹ107000ｹ0, 0000 apple 閨0ｹ ｹ0 6ｦ ｷ 閨0ｹ ｸ70208apple030ｷ302000ｹ107000ｹ0: ｸ ｧ90709 (haemagglutinin-ha) ｸ (neuraminidase-na). 09 閨 ｧ 閨04080ｸ ｸ70208apple ｧ ｦ0apple ｸ ｧ907090ｹ0 (H1 閨0ｷ30ｹ0 H16) ｧ ｦ0apple ｸ ｹ0 (N1 閨0ｷ30ｹ0 039). 020ｸ ｧ ｧ8 fi030ｷ ｷ307 0ｸ7apple08026ｦ0apple0ｷ ｸ ｦ0 ﾞ 02090ｹ ｧ9apple090ｹ0 006ｧ ｸ7 03fi ｸ , apple086ｧ9 閨04080ｸ707 apple ｷ303006ｧ9 fi ｦ0apple0806 HA NA. 6ﾒ ｸ ｦ007, apple ｸ apple , 00 閨040ｷ3 020ｷ apple ｷ ｸ70ｹ0, 閨 ｸ ｦ ｹ ｧ9apple090ｹ0. 6ﾒ ｦ003080ｹ0 020ｷ ｷ ｹ ｧ9apple090ｹ0, fi0ｹ0 applefi 02 0ｸ7 03fi ｸ , apple apple ｷ ｧ9-06, 閨07 006ｧ ｷ , 02 06appleapple ｧ8, ｧ 閨 ｷ3apple08. 6ﾒ2080ｸ ｸ7applefi020ｸ7apple ｸ ｦ ｹ ｧ9apple090ｹ0 ﾞ ｦ ｷ3apple08: 0806 ﾇ ﾇ ｧ ｸ7apple006ｦ0030ｸ ｹ ｹ ｧ9apple090ｹ fi0708, ｧ fi020902, ｹ0 3090ｹ ｧ9 0ｹ0. 6ｾ 閨0409 ﾇ1031, ﾇ ﾇ ｦ ｸ70ｹ ｧ903080ｸ70ｹ0, ﾑ80ｸ70306apple ﾞ ｧ ｧ ｧ ｸ7applefi020ｸ7apple ｸ ｦ ｹ ｧ9apple090ｹ ﾇ ﾇ ｧ ｹ0 040ｸ ｧ90ｹ ｸ70ｹ ｧ ｸ70ｹ ｸ apple080ｸ7, ｹ0 fi000ｷ30ｹ0 apple000306apple ｹ ｹ ｹ ｧ9apple090ｹ ｸ ｧ903080ｸ apple080ｸ70ｹ0 閨04080ｸ ｹ0 00apple ｷ303006ｧ9. 6ﾑ ｧ903080ｸ ｸ70000 fi 閨0ｹ ｹ0 apple080ｸ7 閨04080ｸ ｧ ｸ70ｹ0, fi08080ｸ70ｹ ｹ 閨 閨0ｹ ｸ70ｹ0 閨04080ｸ ｧ apple 03080ｸ7046ｧ ｷ ｷ ｷ ｸ 閨070ｷ , 040ｷ3036ｧ90ｹ fi04080ｸ7. ﾌ ｧ ｹ ｧ9apple090ｹ ｷ3036ｧ9ｨB08- ABSTRACT. The huge epizootics of highly pathogenic avian influenza (subtype H5N1) in Southeastern Asia over the last two years and especially the transmission of avian influenza viruses to humans have alerted the international scientific community. Many support that the threat of a new influenza pandemic appears greater today than ever before. During the 20th century, humanity has faced three pandemics, including the "Spanish flu" of , which claimed over 20 to 40 million lives, and two less dramatic pandemics in and Influenza A viruses are single stranded RNA viruses belonging to the family Orthomyxoviridae. Their genome expresses only 10 proteins, most important of which are the two surface glycoproteins: haemagglutinin (HA) and neuraminidase (NA). So far, 16 different types of haemagglutinin (H1 to H16) and 9 of neuraminidase (N1 to N9) have been recognized. Influenza A viruses are grouped into "subtypes", according to the HA and NA surface proteins they bear (for example H1N1, H5N2). Natural reservoirs of influenza A viruses are the wild aquatic birds (migratory waterfowl), from which all types of HA and NA have been isolated. It is important to mention that migratory waterfowl do not show clinical signs of disease, but shed the virus through their excretions. The host range of flu viruses includes domestic poultry, and mammalian species from aquatic mammals to horses, humans and swine. Because of their segmented single stranded RNA genome, influenza viruses have a very high mutation rate (genetic drift) and the possibility to undergo reassortment. Reassortment may occur when more than one virus co-infect the same cell, exchange genes and as a result, provide a totally new influenza virus (genetic shift). At least two subtypes of influenza A viruses are currently endemic within the human population (H1N1 and H3N2), causing every year outbreaks of disease with very low mortality, especially in elders. Unlike these endemic viruses, pandemic viruses have a much higher morbidity, affecting people of all ages. H1N1, H3N2 and H1N2 influenza viruses are currently circulating in the European and American swine population. Some of the swine influenza virus subtypes, namely H1N1 and H3N2, are thus similar to those of humans, but there are still important antigenic differences between them. Only rarely swine influenza viruses may be transmitted or cause disease to humans. Unlike mammalian influenza viruses, influenza viruses of domestic birds are grouped in two "pathotypes": 6ﾑ ｧ ｦﾐ ｧ9 0ｹ0, ﾒ ｧ8 6ｾ apple ｧ9080ｸ7 縺 ｹ0, Saliburylaan 133, 9820, Merelbeke, Belgium H ｧ9 0ｸ7apple ｧ80ｹ0: H ｧ ｧ904000ｷ30ｹ0: Laboratory of Virology, Faculty of Veterinary Medicine, Ghent University Salisburylaan 133, 9820 Merelbeke, Belgium Submission date: Approval date:

2 13340 ﾒ. 6ｾ409. ﾒ0806ｦﾐ ﾞ ﾒ ﾇ6ｾ4, VAN REETH K ｦ ｧ9000ｹ0: ( ) 02080ｸ70ｹ ｦ00ｹ ｧ80ｹ fi07080ｸ fi ｹ0 (low pathogenic avian influenza viruses LPAI), apple080ｸ7 apple ｦ ｹ ｧ80ｹ0 0208apple06026ｧ ｧ9000ｷ (6 5) 02080ｸ70ｹ ｦ00ｹ0 0ｸ ｧ80ｹ fi07080ｸ fi ｹ0 (high pathogenic avian influenza viruses HPAI), apple080ｸ7 apple ｦ ｧ ｧ8 07fi fi apple080ｸ7 040ｸ % (apple ｷ307 apple fowl plague). 6ｾ402080ｸ70ｹ ｦ00ｹ0 0ｸ ｧ80ｹ fi07080ｸ fi ｹ0 076ｧ802080ｸ707 00fi 閨 閨0409 apple080ｸ7 08 閨03080ｸ ｸ ｧ90709 ﾇ5 6ｧ8 ﾇ7. 09 閨 apple03fi ｸ7036ｧ90ｷ30ｹ0 00 閨 ｸ702 6ｧ9 00apple06ｨB0ｷ308026ｧ ｧ9apple090ｹ0 020ｷ307 apple ﾞ046ｧ , 046ｦ000080ｷ ｧ ｷ3036ｧ ｸ703096ｧ9 0007fi0ｹ ｦ ｦ0 07 apple 閨 apple ｧ ｷ3apple08, 6ｧ apple 03 6ｧ apple 03080ｸ7046ｧ9 0007fi0ｹ ｸ ｧ8 apple080ｸ7 03 apple ｦ ｧ ｧ 閨040ｷ ｸ7 08apple086ｧ9080ｸ 閨080ｹ0 06fi0ｹ0 03 apple ｦ00400 applefi ｷ3apple08. ﾌ06 apple fi ｧ9 040ｸ700080ｷ307086ｦ apple0ｷ30ｹ0 0ｸ702fi fi apple ｦ apple 6ｧ ｧ903080ｹ applefi apple 閨07, fiapple0ｷ30ｹ ｸ707 0ｸ7 0400fi0ｹ ｷ307 apple ｸ apple080ｸ ｧ903080ｸ70ｹ0, 006ｧ ｧ ｸ700apple 閨 apple0ｷ30ｹ ｧ9 020ｷ307 apple ｸ ｸ ｧ ｸ ｧ ｸ ｧ903080ｸ fi apple apple 閨07 applefi 02060ｹ0 00apple06ｨB0ｷ ｧ9000ｹ ｹ ｧ9apple090ｹ0 020ｷ307 apple apple0ｷ3 ﾞ ｧ8 (H5N1) ﾌ ｧ9 (H7N7) ｧ apple0ｷ30ｹ 鐶pple080ｸ apple 03 - apple07070ｷ3 0ｸ7applefi ﾇ apple ｯ 000ｸ703006ｧ9 0ｹ ｷ307 apple apple080ｸ70ｹ apple 03080ｸ7046ｧ9 020ｷ ｸ7apple 閨0ｷ ｹ ｸ ｧ907090ｹ ｹ ｧ9apple090ｹ0 020ｷ307 apple apple06apple00080ｸ702fi02 0ｸ700 閨 ｷ302fi 00apple06036ｧ ｸ ｸ7 07 apple07000ｸ ｦ0 040ｸ704026ｧ ｹ ｸ apple080ｸ7, apple ｦ0080ｸ707 apple0ｷ30ｹ0 08 6ｧ ｹ ｷ3apple080ｹ0 00apple ｧ9 07 apple 閨 ｧ ｧ apple0ｷ apple ﾑ8apple06apple03 閨0807, 閨0ｹ 閨02000ｹ0 apple080ｸ7 閨 陦 apple0ｷ30ｹ0 00fi03060ｹ ,25% 020ｷ ｧ9030ｷ307 apple080ｸ 閨 ｹ0 apple 閨0ｹ0 apple080ｸ apple ｧ9apple09 020ｷ307 apple ｸ702 6ｧ fi ｦ , 閨07 0ｹ ｧ903080ｹ fi 閨0ｹ ｦ ｹ ｧ9030ｷ307, ｦ00ｹ ｧ9apple090ｹ0 020ｷ307 apple ｸ ｧ80ｹ fi07080ｸ fi ｹ0 ( ﾇ ﾇ7037), apple 陦 apple0ｷ30ｹ ｸ 閨 ｧ9, apple apple ｨB ｸ70ｹ0 apple07006ｦ ｹ ｸ ｧ903080ｸ7, 0007 apple ｦ ｧ8 07fi ｧ applefi 04086ｧ ｧ 閨 , fi apple ｦ ｹ ｧ9apple090ｹ0 020ｷ307 apple ( ﾇ5031) apple080ｸ70ｹ Hong Kong, 09 apple 000ｸ703006ｧ9 0ｹ ｧ9apple090ｹ0 applefi ｷ3apple ｷ303006ｧ fi07 6ｦ ｧ ｸ apple ｷ300 閨07 apple ｧ ｷ ｧ ｸ704 6ｧ ﾑ86ｧ fi apple0ｷ30ｹ0 applefi 閨0ｷ30ｹ ｧ apple ｷ303006ｧ9 apple ｹ apple080ｸ7 applefi 06086ｦ00ｹ0 020ｷ307 apple ﾞapple 閨0ｷ30ｹ0 046ｧ 閨 apple ｷ303006ｧ ｹ ｸ apple0ｷ307 applefi 06086ｦ00ｹ ｹ ｧ9apple090ｹ0 020ｷ307 apple (81 applefi fi 026ｦ0apple080ｸ7 ﾇ apple0ｷ3 ﾞ ｧ ｧ applefi ﾇ ﾌ ｧ9 ). ﾇ 040ｸ ｧ80ｹ0 apple ｦ ｹ0 00apple06ｨB0ｷ ｧ80ｹ fi07 0ｹ ｹ ｧ9apple090ｹ0 020ｷ307 apple , fi ｧ 閨0409 0ｸ ｧ80ｹ fi07080ｸ fi ｹ ｸ ｦ0, 08 03fi03080ｹ0 020ｷ307 0ｸ7 03fi6 5060ｷ ｸ apple , 閨080ｹ0 03fi03080ｹ ｸ ｧ903080ｸ ｸ7036ｧ90ｷ30ｹ0 09 apple apple6ｧ8 006ｧ9 0ｹ0 07 陦 0ｹ0 apple ｧ9 0ｹ ｷ3apple08 apple ｦ007 apple03fi ｧ8, ｧ fi020003a ｸ ｧ8 閨03000ｸ707, apple020ｸ 閨0ｷ fi 0ｷ 閨 ｸ ｹ0 apple03 閨 apple06026ｧ907 0ｸ707090ｹ0 ｨB0ｷ30807fi04080ｸ7. low pathogenic avian influenza (LPAI) viruses, which cause localized infections and remain mild or subclinical, and highly pathogenic avian influenza (HPAI) viruses, which cause severe general infection with mortality up to 100% (fowl plague). The majority of avian influenza viruses are low pathogenic and only some, but not all, viruses of H5 and H7 subtypes are highly pathogenic. Occasionally low pathogenic H5 or H7 viruses from wild birds transmit to poultry. Such viruses can undergo mutations in poultry as a result of which they may acquire a highly pathogenic phenotype. Until the recent avian influenza epizootics in Asia, the predominant theory for the creation of a pandemic virus supported that the pig was likely to act as an intermediate host for transmission of influenza viruses from birds to humans. The fact that genetic reassortment between human and avian viruses has also been shown to occur in pigs in nature, had led to the hypothesis that the pandemic viruses of 1957 and 1968 may have been generated through the pig. More recent data, however, come to question these theories and hypotheses: (a) the direct transmission of the H5N1 and H7N7 avian influenza viruses from birds to humans in Southeastern Asia and The Netherlands, and (b) the presence of cellular receptors recognized preferentially by the haemagglutinin of avian influenza viruses in the human conjunctiva and ciliated respiratory epithelial cells, which support that avian influenza viruses can be transmitted in toto (without reassortment) to and between humans or that humans can be the mixing vessel themselves. Furthermore, there is no solid scientific evidence to prove that any influenza virus reassortants, that have originated in swine, have posed a risk for humans. There are three criteria (conditions) an influenza virus must fulfill in order to be characterized as a pandemic virus: (a) it must be a new virus against which humans are immunologically naive, (b) it must be able to replicate in humans causing severe disease, and (c) it must be efficiently transmitted among humans, causing wide outbreaks. So far the H5N1 influenza virus only fulfills the first and second condition, and even though it has been sporadically infecting humans for over two years, it has not yet been able to fully adapt to it ｯs new host. Compared to the human population that may have been exposed to the H5N1 influenza virus in Asia, the number of patients and fatalities due to the H5N1 virus is very small. So far, it appears that swine do not play an important role in the epidemiology of this specific virus. Experimental infections of swine with highly pathogenic H5N1 virus have shown that it does not replicate extensively in pigs. Additionally, extensive serological investigations in the swine population of Viet Nam, indicated that the H5N1 virus merely spread to a very small number ( 0.25%) of contact animals within the epizootic regions. Nevertheless, it is critical to continue monitoring pigs and studying the behavior and spread of influenza viruses in these species. Key words: Avian Influenza, poultry, the role of the pig, ecology, human health implications. 閨 ｹ0 000ｸ ｧ ｹ0: 縺036ｧ9apple09 020ｷ307 apple , apple ｧ9, 08 03fi03080ｹ ｸ ｧ903080ｸ7, ｧ9, 00apple06apple ｹ fi0406 0ｸ709006ｧ9.

3 13KYRIAKIS C. S., VAN REETH K ﾑ8ｦﾐ6ｾ4 ﾞ 縺03 罍ﾇ ﾇ apple06ｨB0ｷ308026ｧ ｹ ｧ9apple090ｹ0 020ｷ307 apple ｧ8 ﾞ046ｧ9 (6ﾒ , , ﾒ 00applefiｨB09, ｦﾐ ｧ9 02..) ｸ702 6ｧ9 6ｦ fi0706, ｸ7036ｧ90ｷ30ｹ ｦ ｧ9apple090ｹ0 020ｷ307 apple apple080ｸ70ｹ0, 閨04080ｸ ｧ 閨 ｹ0 apple 0902fi ｹ0 00apple ｧ80ｹ fi ｹ0, ｸ70ｹ0 08fi6 5080ｸ70ｹ ｹ ｷ3applefi ｹ ｧ9 07 陦 apple ｧ ｧ9apple090ｹ0, apple ｸ ｸ ﾒ , 006ｧ apple ｧ9000ｹ ｧ9apple090ｹ0: , (apple6ｧ ｹ0 1). 6ｾ46ｦ ｷ ｧ ｹ0, 09 apple03 閨 fi03080ｹ0 00ｦﾐ04apple ｧ ｧ9apple ｸ fi020ｷ30400 apple fi ｸ70ｹ0 applefi ｦ apple080ｸ70ｹ0 04 ｯfi fi040008, fi ｦ00306 閨0ｷ30ｹ ｹ ｷ3apple ｾ ｧ908 0ｸ702fi apple03 鐶pple ｧ9 apple0ｷ30ｹ ｹ0 apple ｧ ｹ0, 02fi fi 020ｷ apple0ｷ307 apple080ｸ7 apple 閨 fi ｸ apple080ｸ7 07fi , ｧ apple 03026ｧ8 00apple 閨07 apple080ｸ ｦﾐ04apple ｧ ｧ9apple ｸ fi 閨0ｹ0 0ｸ7apple6ｧ applefi ｹ0 apple ｧ9000ｹ0: ﾞ ｧ ｧ9apple ｧ9apple ｸ7 Hong Kong apple00036ｧ9apple080ｸ ｦ ｦ000 02, 07026ｧ ,2. ﾇ ｸ702 6ｧ9 00apple06ｨB0ｷ308026ｧ ｹ ｧ9apple090ｹ0 020ｷ307 apple ( ｹ0 ﾇ5031) 閨 ｧ8 09 apple ｧ ｧ ｹ ｹ0 apple080ｸ7 閨 apple036ｧ ,4. ﾞapplefi 閨 ｸ ｧ ( ｧ9080ｸ7 2005) 02080ｸ ｷ3apple0806 閨04080ｸ ｧ 閨04080ｸ707 apple (apple apple080ｸ7 閨04080ｸ707 00apple ｷ303006ｧ ｹ0 applefi fi ﾌ fi ｧ9 0ｹ0 (09. ﾌ.08.) (apple6ｧ ｹ0 2) fi applefi 閨04080ｸ707 apple applefi fi0408 6ｧ8 閨04080ｸ ｷ303006ｧ9 apple apple ,3,4. ﾇ ｨB ｧ9 020ｷ ｷ apple ｸ70400fi 02090ｹ0 apple ｧ80ｹ0 07 閨 ｹ ｦ ｸ70306, ｧ90ｹ0 0ｸ7applefi ｸ fi 閨 apple080ｸ7 apple03fi ｷ3apple6ｧ904080ｸ ｷ304086ｧ ｹ0 apple ｧ80ｹ , 08 6ｧ fi fi0ｹ ｧ ｹ0 apple ｧ80ｹ apple ｧ9 07 apple 閨 閨000ｹ0 00apple06ｨB0ｷ308026ｧ9000ｹ01,3,6. ｦﾐ ｧ9 - ﾌ ｧ9-6ﾑ8apple ｧ9 ﾌ ｧ ｹ ｧ9apple090ｹ0 076ｧ802080ｸ 閨 ｷ307 ﾌ ｸ80607 (Orthomyxoviridae), ｷ ｧ ｸ70ｹ0 006ｧ 閨04080ｸ nm fi ｦ00ｹ 閨 fi02030ｷ30708 RNA ( ｸ ｸ802fi 08016ｦ0), ｧ80ｹ0 apple fi ｹ0, ｸ apple086ｧ9080ｹ0 apple ｧ applefi 0306apple ｧ8. 08apple ｸ ｹ ｧ9 026ｦ0apple ｸ ｦ ｹ ｧ9apple090ｹ0: 08 ﾞ, C. ﾌ ｧ ｸ7 026ｦ0apple080ｸ7 C ｧ fi ｷ3apple apple ｦ007 apple03fi fi0406 0ｸ709006ｧ9. ﾌ ｧ9 026ｦ0apple080ｸ7 08 apple ｦ007 04apple ｦ ｹ0 07 apple07000ｸ ｧ80ｹ ｧ9000ｷ301090ｹ ｯ 0ｸ702fi apple 閨07 0ｹ0 06fi0ｹ0 08 apple ｧ apple apple0602fi fi ｾ fi ｧ ｹ ｧ9apple090ｹ0 006ｧ ｸ702086ｧ9 apple080ｸ7 076ｧ802080ｸ ｦ0apple08 ﾞ ｧ9ｨB : ( ) applefi ｦ003080ｹ apple080ｸ7 apple ｸ707, applefi ｷ3apple08 閨0ｷ30ｹ ｧ ｸ7 03fi6 506, (6 5) applefi ｸ ｧ fi ｸ70ｹ ｸ707 0ｸ707ｨB ﾒ2 096ｧ ｹ0 1. ﾌ06 apple ｧ9000ｹ ｧ9apple090ｹ ｸ ｦ : Table 1. Influenza pandemics of the 20th century: ｹ0 6ｾ402 閨 ｹ ｦ0 056ｦ ｦﾐ04apple ｧ ｧ9appleapple ﾇ1031 > ｦ00306 * ﾞ ｧ ｧ9appleapple ﾇ ｦ00306 縺036ｧ9apple ｸ7 Hong Kong ﾇ ｦ * ﾌ ｧ80ｹ fi0ｹ0 020ｷ ｸ ｷ ｹ0 00ｦﾐ04apple ｧ80ｹ ｧ9apple090ｹ ｧ ｷ30402fi0ｹ0, ｧ apple ｧ9 apple0802 閨 apple ｸ apple080ｸ7 閨 apple fi apple ｧ ｧ8: Nicholson K.G., R.G. Webster, A.J. Hay, Textbook of influenza, Blackwell Science, ｧ ｹ0 2. ﾒ03086ｦ ｸ ｦ ｹ ｧ9apple090ｹ0 020ｷ307 apple ﾇ ｧ8 ﾞ046ｧ9, 00apple ｷ300 閨07 applefi ｧ ｸ fi ｸ7 ﾌ ｦ ｧ9 0ｹ0*: Table 2. Cumulative number of confirmed human cases of avian influenza (H5N1) in Southeastern Asia reported to the WHO: ｧ ｹ0 ｦﾐ ｧ ﾒ ﾒ 00applefiｨB09 ﾒ6ｧ907 6ｾ46ｦ ｧ90ｹ ｧ ｧ90ｹ ｧ ｧ90ｹ ｧ ｧ90ｹ ｧ ｧ90ｹ ｧ ｧ90ｹ ｧ ｾ46ｦ * 6ﾒ ｸ702 6ｧ 閨030ｷ30409 applefi ｧ ｸ7 09. ﾌ.08.

4 13342 ﾒ. 6ｾ409. ﾒ0806ｦﾐ ﾞ ﾒ ﾇ6ｾ4, VAN REETH K RNA 02080ｸ ｦ0 020ｷ apple ｦ apple030ｷ ｹ107000ｹ0, 0000 apple 閨0ｹ ｹ0 6ｦ ｸ70208apple030ｷ302000ｹ107000ｹ0 00apple ｧ9 0ｹ0: ｸ ｧ90709 (haemagglutinin-ha) ｸ (neuraminidase-na) (000602fi07 1). ﾇ ｸ ｧ ｸ apple 03080ｸ ｨB080ｸ ｧ 閨 ｧ9: ( ) 040ｸ ｸ ｧ 閨 ｸ ｦ0 applefi ｦ ｸ ｧ (6 5) apple ｦ ｸ70306fi fi07 apple080ｸ ｷ3036ｧ9ｨB applefi apple fi 046ｦ fiapple fi030ｸ ｹ0 6ｧ ｦ0. 09 閨 ｧ , ｦ0apple ｸ ｧ907090ｹ0 ( ﾇ1 0ｷ30ｹ0 ﾇ16) ｦ0apple ｸ ｹ0 (031 0ｷ30ｹ0 039) 閨04080ｸ707 apple ｷ303006ｧ9 (apple6ｧ ｹ0 3) 2,8. 6ｾ46ｦ000080ｷ ﾌ.08., ｧ9 020ｷ ｹ ｧ9appleapple090ｹ0 apple : ( ) ｦ0apple ｸ ｦ0 (A, B 6ｧ8 C), (6 5) ｧ8 applefi apple086ｧ908 apple fi0ｹ0 (000202fi0ｹ0 applefi ｷ3apple08), (09) ｷ ｧ8 apple ｸ70409, ( ) fi 02080ｸ ｦ (00) 0208 閨02080ｹ ｹ0 apple08-00fi070ｷ304090ｹ apple03fi fi 026ｦ0apple080ｸ7 ﾞ, fi0ｹ0/026ｦ0apple080ｹ ｹ ｸ ｧ907090ｹ ｹ ｸ ｹ0 apple080ｸ7 08 閨 ｸ703086ｦ apple 03 閨 , 0906 apple 閨07 0ｹ0 06fi0ｹ0 026ｦ0apple080ｸ7 ﾞ, apple080ｸ7 apple applefi ｷ3apple Hong Kong fi 1, ｨB : ﾞ/Hong Kong/1/82 ( ﾇ3032), 閨07 0ｹ0 06fi0ｹ0 026ｦ0apple080ｸ7 ﾞ, apple080ｸ7 apple applefi 04086ｧ 縺 fi 14, ｨB ﾞ/Swine/Gent/14/2005 (H1N2). 6ﾒ2 0ｸ7 03fi ｸ apple ｦ ｸ ｧ ｧ8 fi030ｷ ｷ307 0ｸ7apple08026ｦ0apple0ｷ ｸ ｦ0 ﾞ 02090ｹ ｧ9apple090ｹ02,7,8. 6ﾒ2 apple fi ｧ9ｨB080ｸ707 08apple ｧ8apple ｧ80ｹ0 07fi0408, apple ｸ fi fiapple ｸ70ｹ0. 6ｾ46ｦ000080ｷ ｸ 閨0ｹ ｦ ｹ0, ｧ9 apple080ｸ ｦ ｯ 0ｸ apple ｦ007 07fi0408 閨04080ｸ ｧ ｧ8 040ｸ7-6ﾑ80602fi07 1. ﾞ07 apple ｷ ｧ9080ｸ ｸ ｦ ｹ ｧ9apple090ｹ0, fiapple080ｸ ｷ3036ｧ9ｨB080ｸ ｸ70208apple030ｷ302000ｹ107000ｹ0 00apple ｧ9 0ｹ0: ｸ ｧ ｸ (apple09096ｧ8: World Health Organization, Avian Influenza: assessing the pandemic threat, January 2005). Figure 1. Schematic representation of an Influenza virus particle, showing the surface glycoproteins: haemagglutinin and neuraminidase (World Health Organization, Avian Influenza: assessing the pandemic threat, January 2005). 0407fi020902, ｧ ｸ70ｹ ｧ ｸ70ｹ ｦ00ｹ0 020ｷ ｧ8 020ｷ ｧ90ｷ apple ｷ apple ｦ apple0ｷ30ｹ0 fi apple03fi ｧ ｹ ｧ9apple090ｹ0 apple080ｸ7 閨04080ｸ707 apple 閨 ｧ8 apple03fi apple 閨04080ｸ707 apple ｧ9000ｹ 閨 , ｧ ｧ ｸ7 03fi ﾒ ｦ003080ｹ0 020ｷ ｷ ｹ ｧ9apple090ｹ0, fi0ｹ0 applefi 02 0ｸ7 03fi ｸ , apple : ｧ apple ｷ , ( ｹ0, ｹ0), ｧ9appleapple08, ｧ ｷ3apple08 (000602fi07 2) 2. 6ﾑ ｧ9 0ｹ ｹ0 086ｦ004090ｹ ｸ ｹ0 020ｷ ｹ ｧ9- apple090ｹ0 ( 閨 fi02030ｷ30708 RNA), 閨0ｹ0/ 閨0ｹ 閨0ｹ0 apple ｦ ｸ70407fi ｧ ｹ0 3. 6ﾒ26ｦ0apple ｸ ｧ907090ｹ0 ( ﾇ) ｸ ｹ0 (03) apple ｧ9 09 ｨB060ｷ307 閨04080ｸ707 apple ｷ303006ｧ9: Table 3. Types of haemagglutinin (H) and neuraminidase (N) and animal species from which they have been isolated: ﾞ ｸ ｧ ｸ7 閨 apple ｷ303006ｧ ｸ ｸ7 閨 apple ｷ303006ｧ9 ﾇ ｷ3apple080ｹ0, 04086ｧ903080ｹ0, ｷ3apple080ｹ0, 04086ｧ903080ｹ0, ﾇ ｷ3apple080ｹ0, ｷ3apple080ｹ0, 04086ｧ903080ｹ0, ﾇ ｷ3apple080ｹ0, 04086ｧ903080ｹ0, 6ｧ9appleapple080ｹ0, 033 ﾇ4 034 ﾇ5 035 ﾇ6 036 ﾇ7 6ｧ9appleapple080ｹ0, 037 6ｧ9appleapple080ｹ0, ﾇ ｧ9appleapple080ｹ0, ﾇ9 039 ﾇ10 ﾇ11 ﾇ12 ﾇ13 ﾇ14 ﾇ15 ﾇ ｧ8: Nicholson K.G., R.G. Webster, A.J. Hay, Textbook of influenza, Blackwell Science, 1998.

5 13KYRIAKIS C. S., VAN REETH K fi ﾞ0ｸ702 閨0ｹ0 00apple ｧ9 07 apple ｧ9ｨb 陦, 0000 apple0802 閨 ｧ ｧ ｧ ｧ ｹ ｸ ｧ907090ｹ0 6ｧ ｹ ｸ ｹ0 (genetic drift), 6ｧ8 fi ｦ ｧ ｦ007080ｸ ｸ702fi ｧ ｦ , 00apple ｧ ｸ ｧ apple0802 閨 ｸ703096ｧ9 0007fi0ｹ ｹ0 07 閨080ｸ ｦ0 (genetic shift) 7,8. 046ｦ 閨 ｸ ｦ ｹ ｧ9apple090ｹ0 ﾞ ｦ apple 0902fi apple apple ｸ70400fi ( 閨 閨 ｹ0 ﾇ 閨07 ﾇ3032) ｨB6ｧ 閨 fi 026ｦ0apple080ｸ7 08 apple ｧ appleapple0602fi fi (apple080ｸ7 apple ｸ707ｨB ｦ000080ｷ ｹ ｧ9000ｹ ｸ7 09. ﾌ.08.). ﾒ fi0708 apple ｦ ｧ9000ｹ ｧ9apple090ｹ fi apple03 076ｧ apple08036ｦ ｧ fi020902, fi07 apple ｹ ｧ9 0ｹ0, ｹ0 0ｸ ｦ ｦ007080ｸ7 (fiapple0ｷ30ｹ ｧ8). ｦﾐ086ｧ9 apple080ｸ apple fi07 apple apple ｧ9000ｹ0 apple ｸ ｨB080ｸ ｧ90ｷ30ｹ ｧ8 040ｸ700apple ﾑ86ｧ fi apple0ｷ30ｹ ｦﾐ04apple ｧ ｧ9apple ｸ fi apple00036ｧ9apple080ｸ % 02080ｸ7 apple 0902fi ｸ7 apple ｸ ｦ0, apple fi ｦ ｧ ｸ ｧ90ｹ ｧ ｹ ｦ fi03060ｹ % 020ｷ ｸ ｷ307 6ｧ ｷ3 020ｷ ,2,9. 6ｾ402080ｸ70ｹ ｧ903080ｸ70ｹ0, 02080ｸ ﾑ80ｸ70306apple ﾞ ｧ8, ｦ 閨 ｷ307 0ｸ7apple08026ｦ0apple0ｷ307: ﾇ1031, ﾇ ﾇ1032 8, fi0ｹ0 apple0ｷ30ｹ ｧ9 0ｸ702086ｧ9 006ｧ ｹ0 040ｸ ｧ90ｹ ｸ70ｹ ｦ00ｹ ｸ apple080ｸ7, ｹ0 apple000306apple ｹ ｹ ｹ ｧ9- apple090ｹ ｸ ｧ903080ｸ apple080ｸ70ｹ0 閨04080ｸ707 00apple ｷ303006ｧ ﾑ ｧ9ｨB ｷ303006ｧ9 apple0ｷ30ｹ 閨0ｹ ｹ0 apple080ｸ7 閨04080ｸ ｧ ｸ70ｹ0, fi ｸ70ｹ ｹ 閨 閨0ｹ ｸ70ｹ0 閨04080ｸ apple 03080ｸ7046ｧ ｷ ｷ ｷ ｸ 閨070ｷ , 040ｷ3036ｧ90ｹ ｯ fi04080ｸ712. ﾌ ｧ ｹ ｧ9apple090ｹ ｷ3036ｧ9ｨB ｦ ｧ9000ｹ0: ( ) 02080ｸ70ｹ ｦ00ｹ ｧ80ｹ fi07080ｸ fi ｹ0 (low pathogenic avian influenza viruses LPAI), apple080ｸ7 apple ｦ ｹ ｧ80ｹ0 07 apple07000ｸ ｧ8 6ｧ ｧ8 0208apple06026ｧ ｧ9000ｷ (6 5) 02080ｸ70ｹ ｦ00ｹ0 0ｸ ｧ80ｹ fi07080ｸ fi ｹ0 (highly pathogenic avian influenza viruses HPAI), apple080ｸ7 apple ｦ ｸ700 閨 ｧ9000ｷ fi apple080ｸ7 040ｸ % (apple ｷ307 apple fowl plague) 2,8. 6ｾ402080ｸ70ｹ ｦ00ｹ0 0ｸ ｧ80ｹ fi07080ｸ fi ｹ0 076ｧ802080ｸ707 apple 閨0409 apple080ｸ7 08 閨03080ｸ ｸ ｧ90709 ﾇ5 6ｧ8 ﾇ7, 040ｷ3036ｧ90ｹ0 0ｸ702fi ｧ apple0ｷ30ｹ0 fi ｧ ｹ ｧ9apple090ｹ0 020ｷ307 apple ｦ0apple080ｸ7 ﾇ5 6ｧ8 ﾇ7 006ｧ ｸ ｧ80ｹ fi07080ｸ fi ｹ0. 6ﾒ ｸ7 03fi ｧ ｸ70207 apple080ｸ ｧ9 080ｸ 閨0409 0ｸ ｧ80ｹ fi07080ｸ fi ｹ ｧ apple ｷ ﾑ8apple6ｧ904090ｹ0, ｧ apple , 閨0409 ﾇ5 6ｧ8 ﾇ ｧ80ｹ fi07080ｸ fi ｹ ｦ 閨0409 0ｸ ｧ80ｹ fi07080ｸ apple 閨04080ｸ 閨0ｹ0 00apple06ｨB0ｷ308026ｧ9000ｹ013. 6ﾑ80602fi07 2. ﾌ fi0ｹ0 026ｦ ｹ ｹ ｧ9apple090ｹ003, ｧ8 07 apple ｹ ｧ9 0ｹ0 020ｷ ｹ ｧ9apple090ｹ0 ﾞ (apple09096ｧ8: Nicholson K.G., R.G. Webster, A.J. Hay, Textbook of influenza, Blackwell Science, 1998). Figure 2. The "wheel of flu", a schematic representation of the ecology of influenza A viruses (Nicholson K.G., R.G. Webster, A.J. Hay, Textbook of influenza, Blackwell Science, 1998). 09 閨 , fi apple ｦ ｹ ｧ9apple090ｹ0 020ｷ307 apple ( ﾇ5031) apple080ｸ70ｹ Hong Kong, 09 apple ｯ 000ｸ703006ｧ9 0ｹ ｧ9apple090ｹ0 applefi ｷ3apple ｷ303006ｧ fi07 6ｦ , 6ｧ ｸ apple ｷ3-00 閨07 apple ｧ ｷ ｧ ｸ704 6ｧ9 03 (apple6ｧ ｹ0 4). 6ﾑ86ｧ fi apple0ｷ30ｹ0 applefi 閨0ｷ30ｹ ｧ apple ｷ303006ｧ9 apple ｹ apple080ｸ7 applefi 06fi 02090ｹ ｧ9apple090ｹ0 020ｷ307 apple ﾑ80706 applefi 閨0ｷ30ｹ0 046ｧ 閨 apple ｷ303006ｧ ｹ ｸ apple0ｷ307 applefi 06086ｦ00ｹ ｹ ｧ9apple090ｹ0 020ｷ307 apple (81 applefi 06fi 026ｦ0apple080ｸ7 ﾇ apple0ｷ3 ﾞ ｧ 閨07 0ｹ ｧ applefi ﾇ ﾌ ｧ9 ) ﾒ ｧ ｷ3036ｧ fi 閨07 6ｾ46ｦ000080ｷ ｧ ｷ3036ｧ ｸ703096ｧ9 0007fi0ｹ ｦ ｦ0 07 apple 閨 apple ｧ9, 6ｧ apple 03 6ｧ apple 03080ｸ7046ｧ9 0007fi0ｹ ｸ ｧ8 (intermediate host), apple080ｸ7 03 apple 閨 ｧ ｧ90803 (mixing vessel) 00 閨040ｷ ｸ7 08apple086ｧ9080ｸ fi0ｹ0 03 apple applefi ｷ3apple08. ﾌ06 apple fi ｧ9 040ｸ700080ｷ307086ｦ apple0ｷ30ｹ fi0308 0ｸ702fi 03 00apple ｦ apple 6ｧ ｧ903080ｹ0 (000602fi07 3) applefi apple 閨 ｧ apple ｦ00ｹ ｸ7-

6 13344 ﾒ. 6ｾ409. ﾒ0806ｦﾐ ﾞ ﾒ ﾇ6ｾ4, VAN REETH K. 096ｧ ｹ0 4. 6ﾑ8apple ｷ300 閨07 apple ｧ9apple090ｹ0 020ｷ307 apple ｷ3apple08 applefi 閨0ｷ30ｹ0 046ｧ : Table 4. Confirmed human cases of avian influenza from 1959 to date: ﾇ ｧ /apple ｧ8 6ｾ402 閨 ｹ0 ﾞ ｧ90ｹ ｧ ﾇ09 ﾞ ﾇ ﾇ ｧ ﾇ Hong Kong ﾇ ﾒ6ｧ907 ﾇ Hong Kong ﾇ ( ) Hong Kong ﾇ (09 03.) ﾌ ｧ9 ﾇ ( ) Hong Kong ﾇ ﾒ ｹ0 ﾇ ﾞ. ﾞ046ｧ9 ﾇ ｧ8: fi ｹ0 ﾌ fi0ｹ ｧ9 0ｹ0 (09. ﾌ.08.). 6ﾑ80602fi07 3. ﾌ06 apple ｧ9000ｹ ｸ ｸ apple fi0ｹ0 03fi03080ｹ ｸ ｧ903080ｸ7 (apple09096ｧ8: Nicholson K.G., R.G. Webster, A.J. Hay, Textbook of influenza, Blackwell Science, 1998). Figure 3. The pandemics of 1957 and 1968, the possible role of swine in the creation of the pandemic viruses (Nicholson K.G., R.G. Webster, A.J. Hay, Textbook of influenza, Blackwell Science, 1998) 閨0ｹ0 04 ｯ 0ｸ702fi ｸ700apple 閨 ﾒ 02 ｯ 03046ｧ807, 080ｸ 閨0ｹ ｦ ｹ0 apple 陦 apple0ｷ30ｹ ｧ ｹ ｧ9apple090ｹ0 apple080ｸ ｸ ｦ ｸ70ｹ ｧ903080ｸ70ｹ0 閨04080ｸ ｧ9 06 apple080ｸ7 apple0308- 閨 fi0408 applefi 06086ｦ00ｹ0 apple fi applefi apple ｸ70ｹ ｦ00ｹ010,14, ｧ ｸ ｧ903080ｸ ｸ apple080ｸ7 閨04080ｸ ｧ 閨 ｧ80ｹ ｹ ﾑ8apple6ｧ904090ｹ0, ｧ903080ｹ0 08 閨 apple07000ｸ fi 02080ｸ7 046ｦ ｸ70ｹ ｦ00ｹ0 0ｸ7apple ｧ90ｹ0 apple080ｸ ｷ3036ｧ9ｨB ｸ ｧ ｹ ｧ9apple090ｹ ｸ apple080ｸ7 (sialic acid 2,6 linkages), ｷ307 apple (sialic acid 2,3 linkages) 閨0ｹ0 閨03000ｸ707000ｹ0 陦 apple0ｷ30ｹ ｧ903080ｹ0 apple ｧ9 閨 ｸ ｧ fi03080ｸ70ｹ ｸ70ｹ ｦ00ｹ0 020ｷ307 apple ｧ80ｹ ｸ ｧ80ｹ fi07080ｸ fi ｹ016. 6ﾒ2 閨03080ｹ0, 006ｧ90400 apple ｧ ｸ707 0ｸ7 0400fi0ｹ ｷ307 apple ｸ apple080ｸ ｧ903080ｸ70ｹ ｦﾐ02 036ｧ9, 040ｷ3036ｧ90ｹ0 fi000ｷ30ｹ ｸ703096ｧ apple06026ｧ907 0ｸ7070ｷ ｷ3apple apple 03 apple07070ｷ3, ｸ703096ｧ9 020ｷ apple080ｸ7 apple ｹ0 apple ｧ9000ｹ ｸ ( ﾇ2032) ｸ ( ﾇ3032) apple08 fi ｸ ｧ903080ｸ7 (000602fi07 4) fi 閨07 applefi apple 閨0ｹ 閨02000ｹ0, applefi apple03fi apple ｧ ｹ ｧ9apple090ｹ0 020ｷ307 apple apple0ｷ3 ﾞ ｧ8 ( ﾇ5031) ﾌ ｧ9 ( ﾇ7037) , 08 6ｧ apple0ｷ30ｹ ｨB080ｸ 閨0ｷ30ｹ0 046ｧ 閨 apple 閨02080ｸ 陦 00030ｷ fi apple08 006ｧ apple 03080ｸ7046ｧ9 020ｷ ｸ7apple 閨0ｷ ｹ ｸ ｧ907090ｹ ｹ ｧ9apple090ｹ0 020ｷ307 apple (sialic acid 2,3 linkages) apple06apple00080ｸ702fi02 0ｸ700 閨07 18, ｷ302fi 00apple06036ｧ ｸ ｸ apple07000ｸ ｦ0 040ｸ704026ｧ ｹ ｸ apple080ｸ719, apple080ｸ ｧ apple ｯ 000ｸ703006ｧ9 0ｹ ｹ ｧ9apple090ｹ0 020ｷ307 apple apple080ｸ70ｹ020,21, ｸ700apple apple06apple ｦ ｹ0 apple080ｸ7 apple ｧ ｦ00ｹ ｧ90ｹ ﾌ ｧ9 21. ﾞapplefi ｧ apple 閨03, 閨0ｹ 閨02000ｹ0 apple080ｸ7 閨 ﾌ ｧ9 23 陦 apple0ｷ30ｹ0 00fi03060ｹ ,25% 020ｷ ｧ9030ｷ307 apple080ｸ7

7 13KYRIAKIS C. S., VAN REETH K ﾑ80602fi07 4. ﾇ ｧ ｷ3036ｧ ｸ703096ｧ ｦ0 apple ｧ9 0ｹ0, ｧ903080ｹ0 apple ｧ ｧ ｧ ｸ apple080ｸ ｷ307 apple ( ﾒ0ｷ307. 6ｾ4apple. ﾒ0ｸ ｹ K. Van Reeth). Figure 4. The "classical" theory for the creation of a pandemic virus, swine is the "mixing vessel" for reassortment of human and avian viruses (C. S. Kyriakis and K. Van Reeth). 6ﾑ80602fi apple ｧ9 08 6ｧ ｹ ｷ3apple080ｹ0 07 apple 閨 ｧ ｧ ｸ apple080ｸ ｷ307 apple ; ( ﾒ0ｷ307. 6ｾ4apple. ﾒ0ｸ ｹ K. Van Reeth). Figure 5. Can humans play the role of the "mixing vessel" for human and avian influenza viruses? (C. S. Kyriakis and K. Van Reeth) 閨 ｹ0 apple 閨0ｹ0 apple080ｸ7 01 閨04apple ｧ9apple09 020ｷ307 apple ｸ702 6ｧ fi ｦ , ｹ ｧ903080ｹ fi 閨0ｹ ｦ ｹ ｧ9-030ｷ307, ｦ00ｹ ｹ ｧ9apple090ｹ0 020ｷ307 apple ｸ ｧ80ｹ fi07080ｸ fi ｹ0 ( ﾇ ﾇ7037), apple 陦 apple0ｷ30ｹ ｸ 閨 ｧ9, apple apple ｨB ｸ70ｹ0 apple07006ｦ ｹ ｸ ｧ903080ｸ7, 0007 apple ｦ ｧ8 07fi ｸ7036ｧ90ｷ30ｹ ｧ applefi 04086ｧ ｧ ,23. 6ｾ apple00036ｧ9apple020ｷ ｷ ｸ 閨070ｷ ｹ ｧ9apple090ｹ0 020ｷ307 apple , 08 6ｧ apple0ｷ30ｹ0 fi fi ｸ ｧ903080ｸ7 0ｷ30ｹ ｸ ｧ ｧ apple 03 6ｧ , apple0ｷ30ｹ ｨB apple fi0ｹ0 03fi03080ｹ ｸ apple080ｸ7. ﾞ08086ｦ fi0ｹ0 ﾇ ｧ fi0ｹ ｦ ｷ3apple08 040ｷ3036ｧ90ｹ ｸ707 0ｸ ｧ9, 02fi ｧ ｹ ｷ3apple080ｹ0 00apple ｧ9 07 apple 閨 ｧ ｧ apple080ｸ apple (000602fi07 5) 22, 23, 24.

8 13346 ﾒ. 6ｾ409. ﾒ0806ｦﾐ ﾞ ﾒ ﾇ6ｾ4, VAN REETH K. 6ﾒ208 6ｾ402 閨 ｹ0 ﾇ ｧ8 ﾞ046ｧ9, 08 ﾒ6ｧ907 0ｸ707080ｹ ｧ9 0ｹ 閨0ｹ0 6ﾑ8apple06apple ｹ ｧ ｸ7 ﾒ ｷ3026ｧ8 6ﾒ208 apple ｦ ｸ 閨04080ｸ70ｹ0 ﾇ ｷ3apple08 apple ｧ 閨 Hong Kong ｸ ｸ ,6,20. 6ﾒ ｧ fi fi , ｸ apple 閨 ｷ3apple0806. ﾇ 07026ｧ ｷ ｸ7 SAR (Special Autonomous Region) Hong Kong 6ｧ : 0803fi ｹ0 08 apple ｸ7-0400fi0ｹ0 020ｷ307 apple ｹ0 apple ｧ80ｹ0 (apple fi applefi apple07apple06000ｹ0, 046ｧ807000ｹ0, apple086ｦ003000ｹ fiapple080ｸ703 ) ｧ apple0ｷ30ｹ ｧ ｸ7026ｧ8 020ｸ 閨040ｷ fi applefi 陦 apple ｧ ｧ9apple090ｹ apple apple080ｸ7 閨07 0ｹ0 06fi0ｹ ｧ9apple090ｹ0 apple080ｸ7 apple apple fi fi ｸ70ｹ apple080ｸ70ｹ ｸ fi apple fi fi ｹ01, 6, 20, 24. ﾌ 06fi0ｹ ｧ ｸ7 Hong Kong , 6ｦ 閨0ｹ ｸ ｸ7 2003, ｹ0 4 apple000306apple ｹ 閨0ｹ ｹ0 apple ｧ80ｹ apple ｧ904080ｸ apple06ｨB0ｷ308026ｧ 閨0ｹ ｹ ｯfi03 0ｸ70207, ｸ702 6ｧ9 00apple06ｨB0ｷ ｧ fi 閨07 0ｹ ｹ0 0903fi02090ｹ0 apple 閨 ,6. 6ﾒ ｦ ｦ apple06ｨB0ｷ ｸ 閨04080ｸ70ｹ0 ﾇ ｹ ｧ9apple090ｹ0 020ｷ307 apple ｧ applefi ｧ ｹ0 ﾒ0803 陦 0ｹ ﾒ2 0ｸ (040ｷ3036ｧ90ｹ0 00apple ｷ3-00 閨 apple ｦ ) ｧ807000ｹ0 0309fi ( 閨 ｸ7 2003). 6ﾒ ｸ702 6ｧ fi apple06ｨB0ｷ308026ｧ9000ｹ ｸ 閨04080ｸ70ｹ0 ﾇ5031 閨04080ｸ ｧ fi07 04 ｯ0803fi ｧ8 ﾞ046ｧ9, ｦﾐ apple0ｷ3076ｧ9, ﾒ6ｧ907, ｷ304046ｧ ｦ0 1,3. ﾞapplefi 閨 ｸ 閨 閨03080ｹ ｸ7 2005, ﾌ.08. 閨 apple apple ｹ ｸ ｦ0 ﾇ apple080ｸ70ｹ0, applefi 02 08apple086ｧ9 74 6ｧ fi03 1,5. ﾒ 0207 apple0704 apple fi fi0ｹ ｸ ｷ307 apple03 鐶pple ｧ ｦ ｹ0, 閨0708 fi0206 apple apple 閨 ｹ0 0208apple0602 閨0ｹ 閨0ｹ0, applefi 閨 ｸ7apple ｧ80ｹ0 0ｸ7apple ｧ ｹ apple 閨0ｹ0 apple 閨0ｹ ｹ ｧ80ｹ0 ﾞ046ｧ9 0ｹ ｯfi03 0ｸ70207, apple ｦ fi ｧ ｸ apple ｧ9. ﾞ ｧ90ｹ0 0ｸ7applefi ｸ7 apple0ｷ30ｹ ｸ7036ｦ apple ｧ ｹ0 09apple0ｷ3 ﾞ ｧ80ｹ ｦ007 apple fi applefi 2 060ｹ ｷ3apple0806, fi0ｹ0 020ｷ ｸ ｷ ｧ ｸ7apple fi0ｹ0. ﾌ 閨 ｧ ｸ7 040ｸ 閨07080ｸ 閨04080ｸ70ｹ ｹ ｧ9apple090ｹ0 020ｷ307 apple apple080ｸ ｦ ｧ ｷ fi, 閨 ｸ70ｹ ｦ00ｹ0 apple080ｸ7 閨04080ｸ ｧ apple fi07, 006ｧ ｧ ｸ ｧ fi ｹ fi ｸ7, ｦ003080ｹ0 020ｷ apple080ｸ7 閨 apple , 閨07000ｹ ｸ 閨0ｹ fi ｹ fi 02090ｹ0 00ｦﾐ04apple ｧ80ｹ ｧ9apple090ｹ0. 6ﾑ apple fi ｧ ｹ ｧ9apple090ｹ ｸ fi 02080ｸ70ｹ ｧ8 (02 0ｸ7 03fi6 506 ), 閨 ｹ0 ﾇ ｧ ｸ7apple 閨0ｸ7030ｸ ｷ ｸ apple ,24. ﾞ07026ｧ , 08 6ｧ apple0ｷ30ｹ fi0ｹ0 apple fiｨB ｹ fi ｹ0 apple07apple06000ｹ 閨07000ｹ 閨0ｹ0 apple ｧ ｷ3036ｧ90ｹ0 07 apple ｧ9 07fi ﾑ8apple6ｧ904090ｹ0, 0906 apple , 06fi0ｹ ｹ ｧ9apple090ｹ fi fi- 020ｷ ｧ9 1. 6ｾ40ｸ707036ｧ802000ｹ ｧ90ｷ304090ｹ apple0ｷ307, apple ｧ9030ｷ ｧ8 apple ｧ ｹ0 6ﾒ ｹ0 ( ﾞ ｧ908 6ﾑ ｧ9080ｸ7 ｦﾐ ｧ9 0ｹ ｹ0 ﾒ ｧ80ｹ0 6ｾ ｧ80ｹ0 縺 ｹ0, ｧ80ｹ0 Maurice Pensaert). Photo 1. Living conditions of humans, poultry and swine in a rural area of Thailand (Archive of the Laboratory of Virology, Faculty of Veterinary Medicine, Ghent University, Professor Maurice Pensaert). 020ｷ ｧ9 2. ﾞ ｨB0ｷ apple080ｸ Hong Kong ( ﾞ ｧ908 6ﾑ ｧ9080ｸ7 ｦﾐ ｧ9 0ｹ ｹ0 ﾒ ｧ80ｹ0 6ｾ ｧ80ｹ0 縺 ｹ0). Photo 2. Live poultry market in Hong Kong (Archive of the Laboratory of Virology, Faculty of Veterinary Medicine, Ghent University).