Journal of General Virology (2013), 94, 21842190 DOI 10.1099/vir.0.

052985-0

Short Identification and characterization of a novel

Communication paramyxovirus, porcine parainfluenza virus 1, from
deceased pigs
Susanna K. P. Lau,1,2,3,43 Patrick C. Y. Woo,1,2,3,43 Ying Wu,1
Annette Y. P. Wong,1 Beatrice H. L. Wong,1 Candy C. Y. Lau,1
Rachel Y. Y. Fan,1 Jian-Piao Cai,1 Hoi-Wah Tsoi,1 Kwok-Hung Chan1
and Kwok-Yung Yuen1,2,3,4
Correspondence 1
Department of Microbiology, University of Hong Kong, Hong Kong, PR China
Kwok-Yung Yuen 2
Research Centre of Infection and Immunology, University of Hong Kong, Hong Kong, PR China
kyyuen@hkucc.hku.hk
3
State Key Laboratory of Emerging Infectious Diseases, University of Hong Kong, Hong Kong,
PR China
4
Carol Yu Center for Infection, University of Hong Kong, Hong Kong, PR China

We describe the discovery and characterization of a novel paramyxovirus, porcine parainfluenza

Received 6 March 2013
Accepted 26 July 2013
virus 1 (PPIV-1), from swine. The virus was detected in 12 (3.1 %) of 386 nasopharyngeal and
two (0.7 %) of 303 rectal swab samples from 386 deceased pigs by reverse transcription-PCR,
with viral loads of up to 106 copies ml1. Complete genome sequencing and phylogenetic
analysis showed that PPIV-1 represented a novel paramyxovirus within the genus Respirovirus,
being most closely related to human parainfluenza virus 1 (HPIV-1) and Sendai virus (SeV). In
contrast to HPIV-1, PPIV-1 possessed a mRNA editing function in the phosphoprotein gene.
Moreover, PPIV-1 was unique among respiroviruses in having two G residues instead of three to
five G residues following the A6 run at the editing site. Nevertheless, PPIV-1, HPIV-1 and SeV
share common genomic features and may belong to a separate group within the genus
Respirovirus. The presence of PPIV-1 in mainly respiratory samples suggests a possible
association with respiratory disease, similar to HPIV-1 and SeV.

Paramyxoviruses are known to cause systemic, exanthe- These are best exemplified by Hendra virus (HeV) (Selvey
matous, respiratory and neurological diseases in a wide et al., 1995; Young et al., 1996) and Nipah virus (NiV)
variety of animals, including humans. They are enveloped, (Chua et al., 1999, 2000). Fruit bats were subsequently
negative-stranded RNA viruses that are classified into two found to be the natural reservoir for both viruses
subfamilies: Paramyxovirinae and Pneumovirinae. Within (Enserink, 2000; Halpin et al., 2000; Young et al., 1996);
the subfamily Paramyxovirinae, there are currently seven however, human outbreaks due to HeV and NiV have
genera: Aquaparamyxovirus, Avulavirus, Ferlavirus, Heni- occurred as a result of transmission from horses and pigs,
pavirus, Morbillivirus, Respirovirus and Rubulavirus. respectively (Chua et al., 1999; Selvey et al., 1995). A
Among members of the Paramyxovirinae, measles virus number of novel paramyxoviruses have since been
(MeV), mumps virus and human parainfluenza virus 14 identified from various animals and their genomes have
(HPIV-14) are the most well-known human paramyx- been sequenced, including Menangle virus (MenPV)
oviruses that cause outbreaks of respiratory/systemic (Bowden et al., 2001), Tioman virus (Chua et al., 2001),
infections (Lau et al., 2005, 2009; Virtue et al., 2009). Tuhoko virus 13 (ThkPV-13) (Lau et al., 2010) and
Cedar virus (CedPV) (Marsh et al., 2012) from bats,
In the last few decades, a number of novel paramyxoviruses Tupaia paramyxovirus (TupPV) from tree shrew (Tidona
belonging to the subfamily Paramyxovirinae have emerged et al., 1999), Salem virus (SalPV) from horse mononuclear
in animals and/or humans, causing severe illness and death. cells (Renshaw et al., 2000), Mossman virus (MosPV)
(Miller et al., 2003), J virus (JPV) (Jack et al., 2005),
3These authors contributed equally to this paper. Beilong virus (BeiPV) or variant (Li et al., 2006; Woo et al.,
The GenBank/EMBL/DDBJ accession numbers for the nucleotide 2012b) and Tailam virus (TlmPV) (Woo et al., 2011) from
sequences of the genomes of PPIV-1 determined in this study are rodents or rodent cells, Tursiops truncates parainfluenza
JX857409JX857411. virus 1 from dolphins (Nollens et al., 2008), Sunshine virus
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from snakes (Hyndman et al., 2012) and feline morbilli-
virus (FmoPV) from domestic cats (Woo et al., 2012a).
During the process of centralized slaughtering of pigs
carried out to ensure safe pork supply in Hong Kong,
deceased pigs are occasionally encountered without any
obvious or identified cause of death, and are excluded from
further food processing. As pigs are also a known reservoir
for a variety of viruses (Lau et al., 2008, 2011), we therefore
hypothesized that previously unrecognized paramyxo-
viruses may be present in our swine population. A novel
paramyxovirus, belonging to the genus Respirovirus, was
identified, with three complete genome sequences deter-
mined. Based on the results, we propose a novel porcine
paramyxovirus, porcine parainfluenza virus 1 (PPIV-1), in
the genus Respirovirus.
A total of 951 samples from 386 deceased pigs were
collected from a slaughter house in Hong Kong from
September 2008 to June 2012 using procedures described
previously (Lau et al., 2008, 2011). These comprised 386
nasopharyngeal, 303 rectal, 153 blood, 56 lung and 53 liver
samples. Viral RNA was extracted using an EZ1 Virus Mini
kit or RNeasy Mini kit (Qiagen). Paramyxovirus detection
was performed by amplifying a 555 bp fragment of the L
gene of respiroviruses using conserved primers (59-
GACTCATCTACTAACGGNTAYGARA-39 and 59-CACA-
AACATCTTGCTACTWATDATNGT-39) as described pre-
viously (Lau et al., 2010; Woo et al., 2012a). As initial
reverse transcription (RT)-PCR revealed a potential novel
paramyxovirus, all samples were subject to RT-PCR for
PPIV-1 by amplifying a 154 bp fragment of the L gene,
using specific primers (59-TTTTGGGTTCAAGAGATT-
CTT-39 and 59-ACCTTTTGGTCTATGTATAAAGA-39).
Real-time quantitative RT-PCR to detect the L gene of
PPIV-1 was performed using FastStart Universal SYBR
Green Master (Roche), with specific primers (59-
TACAATATATGTGGGTGATCCTTACT-39 and 59-GCC-
TGAATCTTCATGATCTTCTAAA-39) as described pre-
viously (Woo et al., 2012a). Three complete genomes of
PPIV-1, from nasopharyngeal samples S033N, S119N and
S206N, were amplified and sequenced using previously
described strategies (Lau et al., 2010; Woo et al., 2011,
2012a, b). Phylogenetic trees were constructed using the
maximum-likelihood method by PHYML 3.0 (Guindon
et al., 2010) with a substitution model selected by
MODELGENERATOR (Keane et al., 2006).
RT-PCR for a 555 bp fragment of the L gene was positive
in two nasopharyngeal samples. Their sequences possessed
,76 % nucleotide identities to those of known paramyx-
oviruses, suggesting a potentially novel paramyxovirus,
PPIV-1. Subsequent specific RT-PCR for PPIV-1 was
positive in 12 (3.1 %) nasopharyngeal and two (0.7 %)
rectal samples. These sequences formed a distinct cluster
separate from known respiroviruses upon phylogenetic
analysis (data not shown). Quantitative RT-PCR showed
that the viral load ranged from 1.36103 to 2.76106 copies
ml21 in positive samples. Attempts to passage PPIV-1 in
various cell lines were unsuccessful.
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Novel porcine parainfluenza 1 virus
The genome of PPIV-1 contained 15396 bases with a G+C
content of 37.437.5 % and conforms to the rule of six as in
other members of the subfamily Paramyxovirinae with
similar genome organization. The three sequenced gen-
omes (GenBank accession nos JX857409JX857411) shared
91.496.7 % nucleotide identity. Their predicted gene
products showed the highest amino acid identities with
those of members of the genus Respirovirus, especially
Sendai virus (SeV) and HPIV-1 (Table 1). Phylogenetic
trees constructed showed that PPIV-1 was most closely
related to SeV and HPIV-1, forming a distinct subgroup
among respiroviruses (Fig. 1). Based on the present results,
we propose to name the new virus porcine parainfluenza
virus 1 (PPIV-1) within the genus Respirovirus.
The genome of PPIV-1 contained a 12 nt complementary
39 leader and 59 trailer sequence, with the exception of an
AAG substitution at the fifth position of the 59 trailer
sequence (Fig. 2). Similar to other respiroviruses, as well as
henipaviruses and morbilliviruses, PPIV-1 contained
characteristic intergenic regions of GAA (genomic sense)
trinucleotides. PPIV-1, SeV and HPIV-1 are common in
having the trinucleotide AAA at the end of the leader
sequence, the trinucleotide GAA at the beginning of the
trailer sequence and using coding frame 3 in the fusion
(F) and haemagglutininneuraminidase (HN) genes. This
is in contrast to BPIV-3, SPIV-3 and HPIV-3, which
possess the trinucleotide GAA at the end of the leader
sequence and the trinucleotide GUA at the beginning of
the trailer sequence, and use coding frame 2 in F and HN
genes.
Similar to most members of the subfamily Paramyxo-
virinae, PPIV-1 contains a conserved AnGn editing site in
the phosphoprotein (P) gene (Falk et al., 2008; Thomas
et al., 1988). In members of the genus Respirovirus, this
RNA editing site is present in all viral species except HPIV-
1 (Matsuoka et al., 1991). To examine the number of G
insertions at the P mRNA editing site, primers (59-
GAAATAATAATGGAGGAAGCCTGGA-39 and 59-AATA-
CATGGCATACGTTTCGGGAT-39) were used to amplify
and clone a 508 bp fragment from strain S206N and the
number of G insertions was determined by sequence
determination of different clones (Lau et al., 2010; Woo
et al., 2012a). Sixty independent clones were obtained, with
mRNAs encoding three putative proteins identified: P
(mRNA sequence the same as the antigenomic RNA
sequence without a G insertion), V (one-G or four-G
insertion) and W (two-G insertion) proteins that shared
the first 323 aa at the N terminus. Among the 60 clones, 44
(73 %) contained the sequence AAAAAAGG (without a G
insertion) (encoding the P protein), 11 (18 %) contained
AAAAAAGGG (one-G insertion) (encoding the V protein),
four (7 %) contained AAAAAAGGGGGG (four-G inser-
tion) (encoding the V protein) and one (2 %) contained
AAAAAAGGGG (two-G insertion) (encoding the W
protein). Similar to other respiroviruses, except HPIV-1,
the editing site of PPIV-1 contained an A6 run. However,
the number of G residues following (two Gs) and the
2185
2186

S. K. P. Lau and others

Table 1. Pairwise amino acid identities of predicted gene products of PPIV-1 compared with those of other paramyxoviruses

Virus* PPIV-1-S206N PPIV-1-S033N PPIV-1-S119N

N P M F A L N P M F A L N P M F A L

Respirovirus
PPIV-1-S206N 95.2 85 96 91.4 93.8 96.7 97 91.8 98.8 96.6 98.6 98.7
PPIV-1-S033N 95.2 85 96 91.4 93.8 96.7 95.1 84 95.4 91 93.8 96.8
PPIV-1-S119N 97 91.8 98.8 96.6 98.6 98.7 95.1 84 95.4 91 93.8 96.8
SeV 74.5 41.7 76.1 60.5 56.5 77.1 75 42.3 75.9 60.4 56.3 76.6 74.8 41.1 76.4 61.2 56.4 76.9
HPIV-1 72.5 41.2 75.6 61 57.1 76.5 72.6 41.1 75.3 62 57 76 72.5 40.8 75.3 61.7 56.9 76.4
BPIV-3 61.8 30.5 62.7 43.6 45.9 61.9 62.1 29.5 62.4 43.1 45.6 61.7 61.6 62.4 43.2 45.7 61.8
SPIV-3 61.7 28.7 61.8 42.9 46.3 61.6 61.8 28.8 61.5 41.9 46.1 61.6 61.1 28.5 61.5 42.8 46.5 61.5
HPIV-3 59.3 29.3 62.6 43.7 47.1 62.2 59.1 29.9 62.6 42.9 46.9 61.9 59.1 29.5 61.8 43.2 47 62.2
Morbillivirus
FmoPV 24.2 16.8 34.3 27.7 20.6 38.4 25.2 18.1 34.6 26.9 20.8 38.7 25.5 18.1 34.2 27.6 21.2 38.4
MeV 24.6 19.3 35.5 27.6 18.8 39.4 25.8 18.2 35.2 27.5 17.8 39.5 24.6 19.2 35 28 18.8 39.5
Avulavirus
APMV-6 20.9 18.5 23.5 28 21.7 27.8 20.4 20.5 23 27.9 22 27.9 20.2 18.8 23.4 28 21.7 27.8
NDV 23.5 17.7 21.3 26.3 25 28 22.6 16.4 20.5 26.5 26.2 27.3 23.3 17.7 21 24.9 25.2 27.7
Henipavirus
HeV 23.9 17.5 33.3 28.8 23.8 38.1 24.3 18 32.2 28.3 22.3 37.8 23.6 17.7 33.6 28.7 23 37.9
NiV 23.8 19.4 34.2 29.8 23.6 38.7 24.3 18 33.6 29.3 23.4 38.2 23.7 18.4 34.4 29.6 23.4 38.5
Rubulavirus
PorPV 22.7 16.1 21.1 25.5 25 30.4 23 17.4 21 26.4 25.3 30.2 22.8 16 21.3 26.1 26 30.3
MenPV 23.9 17.4 19.8 25.6 18.3 29.4 21.9 19 19.3 25 18.7 29.4 23 17 19.8 25.7 17.9 29.4
Unclassified
Paramyxovirinae
AsaPV 27.5 18 41.7 32.9 37.5 50 28.7 18.9 41.4 33 38.5 49.8 28.3 19.2 41.2 32.8 39 49.6
BeiPV 25.3 18.4 36.6 29.5 20.3 38.6 25 20.4 36.3 29 21.1 38.6 25.1 20.5 36.3 30.5 20.5 38.4
TlmPV 24.8 21.1 35.7 29.3 15.4 38.4 24.8 19 35.4 29.3 15.7 38.6 24.5 21 35.4 29.7 15.6 38.2
FdlPV 25.7 20 36.9 29 36.1 42.1 25.2 19.9 37.5 28.8 35.3 42 25.4 17.7 37.2 28.8 35.9 42.2
JPV 23.8 16.4 37.4 29.6 23.6 38.5 23.8 19.6 37.7 30.1 24.1 38.8 23.6 17 36.6 29.2 23.7 38.7
Journal of General Virology 94

MosPV 26.2 18.8 34.8 27.2 20.2 38.5 26.7 16.9 34.5 25.8 20.5 38.3 26 16.6 34.8 26.7 20.7 38.5
TupPV 24.8 19.1 33.1 29.4 21.9 38.2 25.5 18.3 33.1 27.6 22.4 38 25.1 19.5 32.9 29.7 22.1 38.2
NarPV 25.1 18.6 36 27.9 20.6 37.9 25.5 17.9 35.4 27.9 21.3 37.7 25.2 18.7 36.3 27.8 20.3 37.8

*BPIV, bovine parainfluenza virus; SPIV, swine parainfluenza virus; APMV, avian paramyxovirus; NDV, Newcastle disease virus; PorPV, porcine rubulavirus; AsaPV, Atlantic salmon
paramyxovirus; FdlPV, fer-de-lance virus; NarPV, Nariva virus; see text for other abbreviations.

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 Novel porcine parainfluenza 1 virus

893 PPIV-1-S206N (JX857409)

0.5 921 PPIV-1-S119N (JX857411)
649 PPIV-1-S033N (JX857410)
HPIV-1 (NC_003461)
1000 Respirovirus
942 SeV (NC_001552)
HPIV-3 (NC_001796)
923
1000 SPIV-3 (EU439428)
956 BPIV-3 (NC_002161)
AsaPV (EU156171) Aquaparamyxovirus
FdlPV (NC_005084) Ferlavirus
1000 JPV (NC_007454)
TlmPV (JN689227)
652 1000 BeiPV (NC_007803)
SalPV (JQ697837)
FmoPV (JQ411014)
1000
CDV (NC_001921)
954
616 1000 CeMV (NC_005283) Morbillivirus
691 PPRV (NC_006383)
780 MeV (NC_001498)
440 TupPV (NC_002199)
NiV (NC_002728)
Henipavirus
HeV (NC_001906)
461
MosPV (NC_005339)
642 1000 NarPV (NC_017937)
PorPV (NC_009640)
1000 ThkPV-1 (GU128080)
ThkPV-2 (GU128081) Rubulavirus
806 903
ThkPV-3 (GU128082)
627 MenPV (NC_007620)
1000
992 APMV-5 (GU206351)
952 APMV-6 (NC_003043)
APMV-7 (FJ231524) Avulavirus
943 NDV (NC_002617)
1000 GPMV-SF02 (NC_005036)

Fig. 1. Phylogenetic analysis of nucleocapsid (N) gene sequences of PPIV-1 and other paramyxoviruses. The tree was
constructed by the maximum-likelihood method with bootstrap values calculated from 1000 trees and rooted on the midpoint.
Bar, 0.5 substitutions per site. Abbreviations are used according to the nomenclature of the International Committee on the
Taxonomy of Viruses. CDV, canine distemper virus; CeMV, cetacean morbillivirus; PPRV, peste-des-petits-ruminants virus;
GPMV, goose paramyxovirus; see text for other abbreviations.

sequence preceding (ATTGGT) the A6 run were different proteins of HPIV-3, BPIV-3 and SPIV-3, which possess
from other respiroviruses. In addition to the P, V and W multi-basic protein cleavage, furin recognition sites (R-X-
proteins, the P gene of PPIV-1 also encoded a 204 aa K/R-R) that allow intracellular cleavage in trans-Golgi
putative C protein by alternative translation initiation membranes (Morrison, 2003). It has been suggested that F
similar to other respiroviruses. This putative C protein was proteins with a single-basic cleavage site, as observed in
initiated at the second AUG codon in the P/V/C genes, at HPIV-4, must be cleaved by an extracellular host enzyme
the +2 frame relative to P. SeV and HPIV-1 possess a usually found exclusively in the respiratory tract, rendering
nested set of C proteins, called C9, C, Y1 and Y2, which are infections limited to the respiratory tract (Morrison, 2003).
initiated at four different translation starting sites but share
In this study, a novel porcine paramyxovirus, PPIV-1, was
a common C-terminal end (Fasca & Desmecht, 2007;
discovered, which was phylogenetically most closely related
Newman et al., 2002). Such conserved initiation codons
to SeV and HPIV-1. However, in contrast to HPIV-1,
were not found in PPIV-1 (Fig. 2).
which does not possess a mRNA editing function, the P
Similar to SeV and HPIV-1, the F protein of PPIV-1 gene of PPIV-1 is able to produce P, V and W proteins as a
possessed a single-basic protein cleavage site followed by a result of mRNA editing, and the C protein by alternative
highly conserved fusion peptide. This is in contrast to the F translation initiation. Moreover, PPIV-1 is different from
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S. K. P. Lau and others
(a)
PPIV-1-S206N 3 Leader (genome sense)
PPIV-1-S206N 5 Trailer (antigenome sense)
SeV 3 Leader (genome sense)
SeV 5 Trailer (antigenome sense)
HPIV-1 3 Leader (genome sense)
HPIV-1 5 Trailer (antigenome sense)
BPIV-3 3 Leader (genome sense)
BPIV-3 5 Trailer (antigenome sense)
SPIV-3 3 Leader (genome sense)
SPIV-3 5 Trailer (antigenome sense)
HPIV-3 3 Leader (genome sense)
HPIV-3 5 Leader (genome sense)
(b)
3 Leaders
PPIV-1-S206N
SeV
HPIV-1
BPIV-3
SPIV-3
HPIV-3
other respiroviruses in having two G residues instead of
three to five G residues following the A6 run at the editing
site. These results support the idea that PPIV-1 represents a
novel paramyxovirus within the genus Respirovirus.
PPIV-1 is likely to be associated with respiratory disease in
swine, although its pathogenicity remains to be ascertained.
Paramyxoviruses are known for their potential to cross
species barriers and cause severe disease epidemics or
epizootics in the new hosts. Whilst HeV and NiV are
known to cause encephalitis in humans, most of the
recently identified paramyxoviruses have been isolated
from healthy animals or cell lines, and hence their
pathogenicity is largely unknown. Nevertheless, at least
two cases of human MenPV infection with influenza-like
illness have been demonstrated in close contact with
infected pigs using serological studies (Bowden et al., 2001;
Chant et al., 1998). Recently, a novel morbillivirus,
FmoPV, was isolated from the urine of domestic cats,
which was associated with tubulointerstitial nephritis
(Woo et al., 2012a). This suggests that paramyxoviruses
may cause various diseases in different organs of infected
animals. In the present study, PPIV-1 was detected mainly
in nasopharyngeal swabs of deceased pigs with a positive
detection rate of 3.1 %. Whilst the causes of deaths in the
sampled pigs were unknown, the presence of PPIV-1 in a
significant proportion of nasopharyngeal samples with
high viral loads suggested its possible association with
respiratory disease, similar to its close counterparts, HPIV-
1 and SeV (Karron & Collins, 2007). The absence of PPIV-
1 in the available lung samples could be due to the
relatively small sample size compared with nasopharyrn-
geal samples or the possibility that the viral replication was
limited mainly to the upper respiratory tract. However, it is
well known that porcine viruses, such as swine influenza
viruses and porcine reproductive and respiratory syndrome
virus, may play an important role in respiratory illness in
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Fig. 2. Alignment of genome terminal
sequences of PPIV-1 and other respiroviruses.
Dots indicate identical residues. (a) Alignment
of the 39 leader and 59 trailer sequences. (b)
Alignment of the 39 leader sequences.
swine with secondary infection by bacteria such as
Mycoplasma hyopneumoniae through virusbacteria inter-
action (Fablet et al., 2012; Lau et al., 2008). Further studies
on co-infections, serology and cell culture isolation may
provide better insights into the pathogenicity of PPIV-1.
Under the genus Respirovirus, PPIV-1, HPIV-1 and SeV may
belong to a separate group of viruses, group 1, distinct from
another group, group 3, comprising SPIV-3, BPIV-3 and
HPIV-3. PPIV-1 possessed the highest amino acid identities
to HPIV-1 and SeV in their predicted proteins. Phylogenetic
analysis also showed clustering of the three viruses separate
from group 3 viruses (Fig. 1). It has been shown that HPIV-
1 and its murine counterpart, SeV, share high sequence
homology and antigenic cross-reactivity (Lyn et al., 1991).
Therefore, PPIV-1 may share antigenic cross-reactivity with
HPIV-1 and SeV. The three group 1 viruses also possess
other common genome features distinct from group 3
viruses. These include the presence of an AAG substitution
at the fifth position of the 59 trailer sequence, the
trinucleotide AAA at the end of the leader sequence, the
trinucleotide GAA at the beginning of the trailer sequence,
use of coding frame 3 in the F and HN genes, and a single-
basic protein cleavage site in the F protein. Given that they
may be associated with respiratory tract infections in their
hosts, HPIV-1, PPIV-1 and SeV are likely to be closely
related viruses with similar pathogenicity.
The present results support a diversity of paramyxoviruses
circulating in swine populations. Paramyxoviruses belong-
ing to four of the seven paramyxovirus genera, including
Avulavirus, Henipavirus, Respirovirus and Rubulavirus, have
been reported to infect swine (Chant et al., 1998; Chua
et al., 1999, 2000; Heinen et al., 1998; Ishida & Homma,
1978; Janke et al., 2001; Lipkind et al., 1986; Moreno-Lopez
et al., 1986). For respiroviruses, pigs were involved in the
epizootic infection by SeV, the mouse parainfluenza virus
Journal of General Virology 94

1, in Japan during 19531956, although they were probably
not the natural host (Ishida & Homma, 1978). SPIV-3 was
first isolated from the brains of pigs during an epizootic of
interstitial pneumonia and encephalitis in the USA in the
1980s (Janke et al., 2001). However, it was not until
recently that their complete genome sequences were
characterized, which revealed that these viruses were
actually variants of BPIV-3 and were possibly transmitted
from cattle to swine, but failed to establish an active
enzootic state (Qiao et al., 2010). SPIV-3 was mildly
pathogenic to conventionally reared pigs and a serosurvey
carried out on swine farms showed negative results (Qiao
et al., 2010). Cross-species infection of BPIV-3 has also
been suggested in humans (Ben-Ishai et al., 1980), lambs
(Stevenson & Hore, 1970) and pigs (Qiao et al., 2009,
2010). In contrast to SPIV-3, we believe that PPIV-1
represents a novel respirovirus with swine as its major
reservoir, as positive samples were detected throughout the
study period. Given that cross-species transmission of
various viruses has continued to occur from swine to
human (Woo et al., 2012c), continuous surveillance of
swine is important in predicting the potential of porcine
paramyxoviruses in causing new zoonotic epidemics.
Acknowledgements
We would like to thank Dr W. M. Ko, Secretary of Health, Welfare
and Food; Mr Clement Leung, Dr Gloria Tam and Dr W. K. Au of the
Food and Environmental Hygiene Department, Government of the
Hong Kong Special Administrative Region (HKSAR), for their
facilitation and assistance in specimen collection. We are grateful to
the generous support of Mr Hui Hoy and Mr Hui Ming in the
genomic sequencing platform. This work is partly supported by a
Research Grant Council grant (HKU 770211M); the University Grant
Council; the Strategic Research Theme Fund and University
Development Fund, University of Hong Kong; Shaw Foundation;
the Providence Foundation Limited in memory of the late Dr Lui Hac
Minh; a donation from Ms Eunice Lam and the Consultancy Service
for Enhancing Laboratory Surveillance of Emerging Infectious Disease
for the HKSAR Department of Health.
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