Id |
Subject |
Object |
Predicate |
Lexical cue |
T636 |
0-2 |
Sentence |
denotes |
4. |
T637 |
3-13 |
Sentence |
denotes |
Discussion |
T638 |
14-144 |
Sentence |
denotes |
As of February 2020, influenza viruses have been identified and reported in swine from 53 countries worldwide (Table 1; Figure 3). |
T639 |
145-340 |
Sentence |
denotes |
The influenza viruses have been detected in different sample types including swine sera, nasal, tracheal, oropharyngeal, nasopharyngeal swabs as well as oral fluids collected from the live swine. |
T640 |
341-421 |
Sentence |
denotes |
Nasal and snout wipes, lung homogenates and fecal slurry samples were also used. |
T641 |
422-660 |
Sentence |
denotes |
Additionally, the lung as well as other internal organ tissues (Table 2) obtained from either dead or sacrificed swine have also been used for the detection of IAV symptoms i.e., lesions in lungs, pneumonia, bronchitis, bronchiolitis etc. |
T642 |
661-820 |
Sentence |
denotes |
Various methods have been used for the detection of influenza viruses in swine samples depending on the sample type, sample numbers and objective of the study. |
T643 |
821-1147 |
Sentence |
denotes |
Virus isolation methods, using either MDCK, Caco-2, HRT18, or swine testicle cells or the pathogen free embryonated chicken eggs, although considered the gold standard [335,336,337] have largely been taken over lately by the sequencing approaches which tend to provide a considerably faster identification of the IAV subtypes. |
T644 |
1148-1396 |
Sentence |
denotes |
The additional benefit of sequencing over virus isolation is that the sequences would be useful for analyzing the influenza virus outbreak clusters [338], virus evolution or reassortment [339] using phylogenetic analyses in different gene segments. |
T645 |
1397-1584 |
Sentence |
denotes |
A recent study reported that next-generation sequencing can be useful in the influenza virus diagnosis and for the identification of the novel virulence markers and drug resistance [340]. |
T646 |
1585-1731 |
Sentence |
denotes |
Most of the studies have used real-time RT-PCR with matrix-gene specific oligonucleotide primers and TaqMan probe for IAV detection [308,341,342]. |
T647 |
1732-1846 |
Sentence |
denotes |
The conserved sequences of the matrix-gene specific primers can detect any IAV subtype in the swine samples [343]. |
T648 |
1847-2169 |
Sentence |
denotes |
Most of the studies used subtype-specific real-time RT-PCR for the IAV subtyping, however, few studies opted for the conventional approach of reverse-transcription PCR followed by Sanger sequencing for amplification of the HA and/or NA genes for retrieving the sequences for phylogenetic analyses to identify the subtypes. |
T649 |
2170-2315 |
Sentence |
denotes |
Although the real-time RT-PCR is a powerful and rapid tool for the subtyping of IAV strains, it is more expensive than reverse-transcription PCR. |
T650 |
2316-2610 |
Sentence |
denotes |
A few studies have reported reverse-transcription PCR based amplification of all the eight gene segments of IAV to generate the whole genome sequences [33,91,128,231] but in most cases, the whole genome sequences were generated using MiSeq next-generation sequencing approach [150,214,325,329]. |
T651 |
2611-2738 |
Sentence |
denotes |
A great advantage of this sequencing approach is that it can identify novel influenza viruses in the swine samples [80,90,301]. |
T652 |
2739-2906 |
Sentence |
denotes |
Most of the serological investigations used one or more methods for influenza virus detection and subtyping in the swine samples e.g., ELISA, HI, NI, MN, or VN assays. |
T653 |
2907-3027 |
Sentence |
denotes |
The serological methods are useful in large-scale surveillances for screening large number of samples in a limited time. |
T654 |
3028-3254 |
Sentence |
denotes |
However, the molecular detection assays are more reliable than the serological methods given the higher sensitivity, but the serological assays are rapid and affordable hence are preferred for large-scale surveillance studies. |
T655 |
3255-3406 |
Sentence |
denotes |
The molecular and serological investigations report either active infections (viral RNA) or past exposures (antibodies) in swine samples, respectively. |
T656 |
3407-3573 |
Sentence |
denotes |
The molecular detection approaches followed by sequencing are largely used for the research focusing on the influenza virus epidemiology [73,78,82,87,94,128,146,150]. |
T657 |
3574-3822 |
Sentence |
denotes |
Several studies used histological examinations e.g., immunohistochemistry or immunofluorescence to identify the IAV symptoms in lung or other internal organs of either dead or severely sick swine sacrificed for the investigations [130,189,315,318]. |
T658 |
3823-3950 |
Sentence |
denotes |
Immunohistochemistry provides a rapid and affordable diagnosis of the influenza virus disease using swine tissue samples [344]. |
T659 |
3951-4083 |
Sentence |
denotes |
One major benefit of immunohistochemistry is that it can be used in the retrospective analysis of the archived tissue samples [245]. |
T660 |
4084-4404 |
Sentence |
denotes |
A large number of investigations have reported sub-clinical influenza virus infections in asymptomatic (apparently healthy) swine [78,103,150,161,171,278,300], indicating that influenza infections can go undetected while the swine may be shedding virus and hence may infect other swine and farm workers in contact [183]. |
T661 |
4405-4626 |
Sentence |
denotes |
Intriguingly, most of the swine samples processed in Australia, Europe, and North America were obtained from the symptomatic swine while most of the Chinese swine samples were collected from asymptomatic swine (Figure 5). |
T662 |
4627-4966 |
Sentence |
denotes |
Symptomatic swine may exhibit mild or severe influenza like symptoms [59,202,252], including fever, coughing, sneezing, pneumonia, bronchitis, reduced appetite, diarrhea, nasal and/or ocular discharge, conjunctivitis, weakness, anorexia, prostration, weight loss, abortion in sows, and mortality in some cases [89,224,231,239,248,282,318]. |
T663 |
4967-5079 |
Sentence |
denotes |
Most studies where the swine were severely infected reported reduced appetite and weight loss [129,246,247,292]. |
T664 |
5080-5226 |
Sentence |
denotes |
Due to IAV infection, the swine takes longer to weigh 100 kg body mass [226], hence the IAV disease burden affects the swine farmers economically. |
T665 |
5227-5380 |
Sentence |
denotes |
Varying rates of mortality of swine due to IAV infections were reported from around the world ranging from 0.5% to 30% [138,206,213,218,221,273,296,327]. |
T666 |
5381-5639 |
Sentence |
denotes |
This wide difference in mortality rate could be due to novel virus strains emerged through reassortments within the swine [273,315,327] or inter-species transmission, e.g., avian to swine transmission, resulting into severe disease outbreaks [27,80,119,133]. |
T667 |
5640-5857 |
Sentence |
denotes |
One example of the emergence of a novel influenza virus strain is the emergence of A(H1N1)pdm09 strain due to reassortment between avian and swine IAV strains in swine which resulted into 2009 influenza pandemic [33]. |
T668 |
5858-6087 |
Sentence |
denotes |
Additionally, the emergence of IAV subtype H1N2 is another classic example of influenza virus reassortment which resulted into severe disease outbreaks in Japanese and Korean swine populations during 1990s and 2000s [27,121,140]. |
T669 |
6088-6210 |
Sentence |
denotes |
Strains of IAV can infect the swine of any age group; piglets as young as one week may become infected with IAV naturally. |
T670 |
6211-6476 |
Sentence |
denotes |
Interestingly, a study in Denmark observed a piglet as young as just three days was infected with IAV despite having maternally derived IAV antibodies [183], suggesting that the infection might have occurred from the infected sow which was shedding the virus [183]. |
T671 |
6477-6678 |
Sentence |
denotes |
However, the symptoms of the influenza-like illness in swine may last only for one week but the virus shedding may still persist until 41 days after appearance of the influenza-like symptoms [183,296]. |
T672 |
6679-6852 |
Sentence |
denotes |
This phenomenon may have serious implications in influenza virus spill over to the non-infected swine as well as to the exposed farm workers due to prolonged virus shedding. |
T673 |
6853-7055 |
Sentence |
denotes |
Three other studies observed virus shedding in swine and reported that the virus shedding may persist until the 11 day [256], 20 day [255], or 29 day [296] after onset of the clinical symptoms in swine. |
T674 |
7056-7175 |
Sentence |
denotes |
This variation in the duration of the virus shedding might be strain dependent, which needs to be further investigated. |
T675 |
7176-7427 |
Sentence |
denotes |
A higher rate of virus shedding and IAV prevalence was reported during the fall and early winter months than summer season because the high relative humidity present in the environment during summer decreases the transmission of influenza virus [267]. |
T676 |
7428-7655 |
Sentence |
denotes |
The high relative humidity in summer facilitates the generation of larger droplets which are less likely to be aerosol transmitted to a longer distance as they tend to fall on the ground quickly after their formation [240,258]. |
T677 |
7656-7804 |
Sentence |
denotes |
Several cases of inter-species transmission were identified which documented transmission of IAV between human and swine or between birds and swine. |
T678 |
7805-8146 |
Sentence |
denotes |
The occurrence of the avian influenza virus strains in swine in China (H5N1, H9N2, H4N1, H4N6, H5N3, H10N5, H4N8, H6N6, H7N9), United States (H4N6, H6N2, H7N2), Canada (H4N6, H3N3), South Korea (H7N2, H5N2), Nigeria (H5N1), and Egypt (H5N1, H5N2, H9N2 ) serve as the evidence of interspecies transmission of avian influenza viruses to swine. |
T679 |
8147-8286 |
Sentence |
denotes |
The first evidence of avian influenza virus active infection in swine appeared in 1999 in Canada when H4N6 virus was isolated from a swine. |
T680 |
8287-8414 |
Sentence |
denotes |
Later several other avian-origin IAV strains were detected and sequenced in swine in China, Canada, and South Korea (Figure 6). |
T681 |
8415-8672 |
Sentence |
denotes |
Various studies have spotted wild birds visiting the swine farms or in the vicinity which suggested that wild birds may have served as the carriers for the introduction of the different avian-origin IAV subtypes to the swine populations [59,80,100,246,332]. |
T682 |
8673-8889 |
Sentence |
denotes |
The highest number of avian-origin IAV strains were reported in Chinese swine which shows frequent avian-swine interaction in China, a country that has historically been an epicenter for influenza virus disease [69]. |
T683 |
8890-9002 |
Sentence |
denotes |
Egypt is recognized as a “hot spot” for the influenza virus reassortment due to its geographical location [345]. |
T684 |
9003-9233 |
Sentence |
denotes |
The role of migratory wild birds in the introduction of avian influenza in Egypt has been already established [346,347], and the highly pathogenic strains of the IAV have previously been detected in migratory birds in Egypt [348]. |
T685 |
9234-9531 |
Sentence |
denotes |
Since migratory wild birds were reported to harbor in the vicinity of Cairo [60] therefore, the probability of the migratory bird–swine interaction in the regions remain high which very well explains the occurrence of highly and low pathogenic strains of avian-origin IAV in swine in Cairo, Egypt. |
T686 |
9532-9775 |
Sentence |
denotes |
Given the “mixing vessel” nature of the swine, the occurrence of avian-origin IAV strains in swine is alarming in terms of IAV reassortment and evolution which may trigger the emergence of novel IAV strains of pandemic potential in the future. |
T687 |
9776-10087 |
Sentence |
denotes |
Further, the multiple reports of double or triple reassortant IAV strains in swine are evidence that IAV co-infections may facilitate the antigenic diversity of the influenza viruses; and as a result, new HA and NA subtypes of IAV may be continually added to the existing 18 HA and 11 NA subtypes in the future. |
T688 |
10088-10267 |
Sentence |
denotes |
Intriguingly, the frequency of the occurrence of double or triple-reassortant IAV strains in swine has dramatically increased in the recent decades [76,81,87,109,161,250,268,278]. |
T689 |
10268-10498 |
Sentence |
denotes |
One unique example of the reassortment and evolution of the pandemic strain of IAV in swine was the emergence of A(H1N1)pdm09 virus in swine in Mexico which evolved due to the reassortment between avian and swine IAV strains [33]. |
T690 |
10499-10838 |
Sentence |
denotes |
While an overwhelming majority of investigations reported IAV in the swine across the world (Figure 7A), there were only a few reports which documented either active infections or past exposures of the swine to the influenza virus types IBV [37,171,186,277] (Figure 7B), ICV [38,117,186] (Figure 7C) or IDV [5,43,44,45,46,217] (Figure 7D). |
T691 |
10839-11057 |
Sentence |
denotes |
A low prevalence of IBV was observed in swine given that only one study reported the IBV antibodies in swine samples in England during 1991–1992 [186] with no evidence of further spill over to other European countries. |
T692 |
11058-11161 |
Sentence |
denotes |
The active infection of IBV was later reported in US swine when two IBV isolates were obtained in 2009. |
T693 |
11162-11365 |
Sentence |
denotes |
A recent study from Taiwan reported three strains of the Victoria/B lineage of IBV in naturally infected swine in 2014 [171], again there was no further report of dissemination to nearby Asian countries. |
T694 |
11366-11448 |
Sentence |
denotes |
The IBV infected swine were apparently healthy with no signs of influenza disease. |
T695 |
11449-11631 |
Sentence |
denotes |
The first report of ICV appeared in Chinese swine after the virus was isolated from apparently healthy swine in 1981 in a routine diagnostic procedure at an abattoir in Beijing [38]. |
T696 |
11632-11788 |
Sentence |
denotes |
Later ICV seroprevalence was reported in English and Japanese swine during 1980s–1990s [117,186] with no further evidence of circulation anymore thereafter. |
T697 |
11789-11958 |
Sentence |
denotes |
The IDV was first detected and characterized in 2011 in Oklahoma based swine in the United States which appeared to have made a species jump from cattle to swine [5,43]. |
T698 |
11959-12142 |
Sentence |
denotes |
Interestingly, a complete IDV genome was retrieved from a symptomatic sow in Italy in 2015 which was found closely related to the IDV genome reported in 2011 from Oklahoma, USA [217]. |
T699 |
12143-12244 |
Sentence |
denotes |
This might have happened due to the trade of the cattle or swine between Italy and the United States. |
T700 |
12245-12475 |
Sentence |
denotes |
A recent study from China identified IDV sequences which shared a high similarity (99%–100%) with the IDV sequences reported earlier from the cattle in China [102] which was another evidence of bovine to swine transmission of IDV. |
T701 |
12476-12622 |
Sentence |
denotes |
The IDV has been in circulation in swine in the current decade with reports emerging from swine in Italy, Luxembourg, China and the United States. |
T702 |
12623-12819 |
Sentence |
denotes |
In summary, IAV was first isolated from a swine in USA in 1930 [34,349] and later antibodies for the human influenza viruses were reported in swine at the State Prison of New Jersey in 1937 [259]. |
T703 |
12820-12984 |
Sentence |
denotes |
More IAV outbreaks and cases in swine in North America were reported during 1981–2000; the frequency has now dramatically fallen in the last two decades (Figure 8). |
T704 |
12985-13095 |
Sentence |
denotes |
This might be due to improved swine influenza surveillance and vaccination in North America in recent decades. |
T705 |
13096-13248 |
Sentence |
denotes |
The H1N1, H1N2, H3N2, and A(H1N1)pdm09 viruses were reported from the commercial, backyard, exhibition, feral swine and wild boars in the United States. |
T706 |
13249-13445 |
Sentence |
denotes |
As of February 2020, the highest number of reports of influenza virus infections in swine in a country were documented in the United States (n = 40) followed by China (n = 39) and Canada (n = 24). |
T707 |
13446-13576 |
Sentence |
denotes |
The highest number of IAV positive swine samples were reported in the United States (36128/200384) followed by China (5031/90760). |
T708 |
13577-13723 |
Sentence |
denotes |
One of the factors behind the higher number of IAV cases in swine in the United States compared to China would be related to the disease symptoms. |
T709 |
13724-13967 |
Sentence |
denotes |
A majority of the North American swine samples that were screened for the IAV infections had either mild or severe symptoms of influenza-like illness which would have made it visually easier to identify IAV infected swine in the United States. |
T710 |
13968-14176 |
Sentence |
denotes |
On the contrary, a smaller number of Chinese swine exhibited influenza-like disease symptoms while a large proportion of the Chinese swine population appeared to have sub-clinical infections with no symptoms. |
T711 |
14177-14292 |
Sentence |
denotes |
This would have made it difficult for identifying the influenza virus infected swine during surveillances in China. |
T712 |
14293-14428 |
Sentence |
denotes |
The first report of IDV in 2011 in Oklahoma swine reflected the antigenic diversity and evolution of influenza viruses in the US swine. |
T713 |
14429-14686 |
Sentence |
denotes |
However, the recent influenza virus disease prevalence in North American swine appeared to have declined, nevertheless, given the large swine population of the continent, the surveillance should continue to track the influenza virus diversity and evolution. |
T714 |
14687-14959 |
Sentence |
denotes |
The first serological evidence of IAV in European swine was documented from the Czechoslovakia during 1969–1972, but the first H1N1 virus in European swine was isolated in Belgium in 1979 which was apparently transmitted from wild ducks in Germany to the swine in Belgium. |
T715 |
14960-15113 |
Sentence |
denotes |
Since then several H1N1, H1N2, H3N2, and A(H1N1)pdm09 viruses have been detected in commercial and backyard swine as well as in wild boars within Europe. |
T716 |
15114-15279 |
Sentence |
denotes |
The incidence of IAV in European swine has increased several folds in the past two decades with a relatively high number of IAV positive swine samples (19644/49814). |
T717 |
15280-15389 |
Sentence |
denotes |
Most of the IAV positive European swine were reported having influenza-like symptoms at the time of sampling. |
T718 |
15390-15557 |
Sentence |
denotes |
Germany reported the highest number of IAV positive swine in Europe where the pork industry is considered the third largest globally after China and the United States. |
T719 |
15558-15859 |
Sentence |
denotes |
Importantly, the IDV was more recently identified in the European swine, first in Italy in 2015, and later a retrospective study identified IDV infection in swine samples collected in Luxembourg during 2014–2015 which indicated that the circulation of IDV in European swine took place only after 2014. |
T720 |
15860-15927 |
Sentence |
denotes |
The evidence has suggested the bovine to swine transmission of IDV. |
T721 |
15928-16145 |
Sentence |
denotes |
This observation is interesting because until recently more emphasis has been given to the avian-swine interaction and the bovine–swine interactions have been neglected from the influenza virus spill over perspective. |
T722 |
16146-16291 |
Sentence |
denotes |
The first occurrence of IAV in Asian swine can be traced back to 1969, but the IAV prevalence has increased multi-fold in the recent two decades. |
T723 |
16292-16400 |
Sentence |
denotes |
The IAV subtypes H1N1, H1N2, H3N2, and A(H1N1)pdm09 have become endemic in swine in several Asian countries. |
T724 |
16401-16569 |
Sentence |
denotes |
The highly pathogenic avian-origin IAV strains of H5N1 and H7N9 have been reported from Chinese swine while H5N1 has been documented in swine in Viet Nam and Indonesia. |
T725 |
16570-16649 |
Sentence |
denotes |
The highly pathogenic strains of H5N2 have been reported in South Korean swine. |
T726 |
16650-16765 |
Sentence |
denotes |
Several LPAIV strains including H4N1, H4N6, H4N8, H6N6, H9N2, and H10N5 have also been documented in Chinese swine. |
T727 |
16766-16930 |
Sentence |
denotes |
The studies suggested a frequent interaction between wild birds and swine in China which appeared to have transmitted avian-origin IAV strains in the Chinese swine. |
T728 |
16931-17056 |
Sentence |
denotes |
Occurrence of equine influenza virus H3N8 in Chinese swine further expanded the genetic diversity of swine influenza viruses. |
T729 |
17057-17267 |
Sentence |
denotes |
Despite an avian to human transmission of certain avian influenza virus strains including H5N1 virus, only a limited human to human transmission of avian influenza viruses was established in the past [350,351]. |
T730 |
17268-17555 |
Sentence |
denotes |
With the passage of these avian-origin IAV strains in a mammalian host like swine, a high probability remains of these avian influenza virus strains to adapt and gain the ability of the human to human transmission, if happens, the consequences would be devastating for the public health. |
T731 |
17556-17707 |
Sentence |
denotes |
Australian swine were free from influenza virus until the year 2009 when a New South Wales swine farm reported an influenza-like outbreak in the swine. |
T732 |
17708-17812 |
Sentence |
denotes |
The zoonotic transmission of the A(H1N1)pdm09 virus was reported to the farm workers and the farm owner. |
T733 |
17813-17941 |
Sentence |
denotes |
Until now, there have only been four IAV reports in Australian swine which reflects a low prevalence of IAV in Australian swine. |
T734 |
17942-18056 |
Sentence |
denotes |
New Zealand is yet to officially report the influenza virus prevalence in swine and remains free from the disease. |
T735 |
18057-18231 |
Sentence |
denotes |
A retrospective study identified that the A(H1N1)pdm09 virus was present in Mexican swine as early as 2000, well before the influenza pandemic occurred during March–May 2009. |
T736 |
18232-18450 |
Sentence |
denotes |
A high genetic diversity of IAV in Mexican swine due to live swine imports from North America and Europe during 1980s laid the foundation of the emergence of zoonotic strain of A(H1N1)pdm09 virus in Mexican swine [33]. |
T737 |
18451-18559 |
Sentence |
denotes |
The highest number of IAV positive swine in Central America were reported from Mexico followed by Guatemala. |
T738 |
18560-18725 |
Sentence |
denotes |
Interestingly, a report of the highly pathogenic H5N2 virus in Mexican swine in 2018 further triggers the alarm in the context of a potential novel IAV reassortment. |
T739 |
18726-18881 |
Sentence |
denotes |
Outbreaks of IAV in the South American swine populations occurred during the last two decades, with the highest prevalence reported in the Brazilian swine. |
T740 |
18882-19034 |
Sentence |
denotes |
A considerable proportion of cases showed sub-clinical infections with no symptoms which might have made it more difficult to detect the infected swine. |
T741 |
19035-19153 |
Sentence |
denotes |
The reports of IAV active infections or the seroprevalence appeared in the African swine only during last two decades. |
T742 |
19154-19283 |
Sentence |
denotes |
Until February 2020, IAV have been detected in swine in Cameroon, Nigeria, Egypt, Kenya, Reunion Island, Uganda, Togo, and Ghana. |
T743 |
19284-19487 |
Sentence |
denotes |
However, South Africa has a considerable swine population [352], but currently there is no published report on the prevalence of active IAV or other influenza virus infections in the South African swine. |
T744 |
19488-19626 |
Sentence |
denotes |
This might be because of the lack of an active surveillance for the detection of the influenza virus disease in the swine in South Africa. |