사람 메타뉴모바이러스

사람 메타뉴모바이러스 (HMPV)
생물 분류ℹ️
역: 바이러스
계: 리보바이러스계
문: 네가르나바이러스문
강: 몬지바이러스강
목: 모노네가바이러스목
과: 뉴모바이러스과
속: 메타뉴모바이러스
종: 사람 메타뉴모바이러스

사람 메타뉴모바이러스(human metapneumovirus, hMPV)는 뉴모바이러스과에 속하는 음성 단일가닥 RNA 바이러스이다.[1] 조류 메타뉴모바이러스 C형과 계통학적으로 매우 밀접하다. 2001년에 네덜란드에서 RAP-PCR(RNA Arbitrarily Primed -)기술을 통해 처음으로 분리되었다.[2] 근래에 유아의 호흡기 세포융합 바이러스 감염이 감소함에 따라 현재 유아 급성 호흡기 감염에 있어서 두 번째로 흔한 감염원이다.

hMPV는 2개월에서 3개월 사이의 유아에게서 피크를 찍는 호흡기 세포융합 바이러스보다 살짝 늦은 6개월에서 12개월 사이의 유아에게서 피크를 찍는다. 호흡기 세포융합 바이러스와 hMPV의 위험성과 의료적 증상은 서로 유사하다. 노인 역시 hMPV에 취약한 대상이다.

분류

메타뉴모바이러스속에 속하는 바이러스종과 이름[3]
속 종 바이러스 (축약어) NCBI 분류 ID
메타뉴모바이러스 조류 메타뉴모바이러스* 조류 메타뉴모바이러스(AMPV) 38525
사람 메타뉴모바이러스 사람 메타뉴모바이러스(hMPV) 162145

*모식종

유행병

어린이 입원환자 및 외래환자의 기도 감염 원인에서 hMPV는 6%에서 40%를 차지한다.[4][5][6][7][8][9][10][11][12][13] 전세계적으로 퍼져있으며, 온대지방에서는 늦겨울에서 봄 사이에 호흡기 세포융합 바이러스, 인플루엔자와 함께 계절성 바이러스로 따라온다.[4][14][15] 또한 5세 이하의 어린이의 대부분이 이 바이러스에 노출되어있음이 혈청학 연구로 밝혀졌다.[2][16][17][18][19][20][21] 이처럼 거의 모든 사람이 어린 시절에 이 바이러스에 감염되지만, 이후에 다시 이 바이러스에 재감염되는 일이 빈번하게 일어난다.[4][22][20][23]

인간 메타뉴모바이러스로 인해 야기되는 대표적 질병은 감기(상부 기도 감염)이다. 그러나 미숙아,[24] 65세 이상의 노인,[23][25][26] 면역결핍 환자의[27][28][29][30] 경우 감기보다 더 치명적인 질병이 걸려 입원환자로 전환되는 경우가 있다. 노인에게 hMPV가 인플루엔자만큼 치명적인 바이러스로 작용함이 입원환자와 응급실 방문환자를 대상으로 한 연구를 통해 밝혀졌다.[23][25][26][31] 또한 천식 환자나[32][33][34][35] 만성 폐쇄성 폐질환[36][37][38] 환자의 경우 hMPV 감염은 더욱 치명적일 수 있다. 장기 간호시설의 환자들 사이에서 hMPV가 급증해 사망자가 발생한 사례가 다수 있다.[39][40][41][42][43] 2020년 일본 아와시의 한 노인시설에서는 17명의 hMPV환자가 발생해 5명이 사망했다.[44]

유전체

hMPV의 유전체 구조는 호흡기 세포융합 바이러스와 유사하나 몇 가지 차이점이 존재한다. 첫째로 hMPV에는 비구조유전자 NS1, NS2가 존재하지 않으며, 둘째로 hMPV의 안티센스 RNA 유전체에 있는 8개의 오픈 리딩 프레임이 호흡기 세포융합 바이러스의 3’-N-P-M-F-M2-SH-G-L-5’과는 다르다.[45] 이보다는 조류 메타뉴모바이러스와 훨씬 유사하며, 특히 조류 메타뉴모바이러스 C와 유사하다. 계통학적 분석을 통해 hMPV에 소분류로 A와 B가 존재하며, 각각은 또 A1/A2와 B1/B2의 아류형으로 구성됨이 밝혀졌다.

바이러스학

hMPV는 코와 허파의 상피세포를 감염시키는데, 당단백질을 통해 헤파란황산염 및 글리코사미노글리칸과 반응하여 세포막에 부착할 것으로 추측된다. 또한 hMPV의 융합 단백질에 있는 아르기닌-글라이신-아스파라긴(RGD) 모티프가 인테그린을 세포 수용체삼아[46][47][48][49] 바이러스가 세포 내부에 엔도솜의 형태로 침투할 수 있도록 한다.[50][51]

검출

hMPV는 역전사 중합효소 연쇄반응(RT-PCR)을 통해 호흡기 표본으로부터 RNA를 증폭하여 검출한다. 이외에도 핵산을 이용해서 저렴하게 검출할 수 있는데, 구체적인 방법은 다음과 같다.

  1. 면역 형광법을 통해 비인두(鼻咽頭) 분비물로부터 직접 hMPV 항원을 찾는다.
  2. 단클론항체를 사용한 면역형광염색을 통해 비인두(鼻咽頭) 분비물을 직접 찾거나, 아니면 이를 쉘 바이알(shell vial)에서 배양한 후 찾는다.
  3. 면역형광검정법(- assay)를 통해 hMPV 특이적 항체를 찾는다.
  4. 다클론항체를 통해 배양된 세포로부터 직접 분리한다.

전염

전염 경로가 완전히 밝혀지지는 않았으나, 감염된 분비물, 비말, 에어로졸, 혹은 매개물을 통해 전염될 것으로 여겨진다. 병원에서 hMPV에 감염된 사례도 보고되었다.[52]

치료

아직 뚜렷한 치료법이 개발되지 않았으나,[53] 모형동물에서는 리바비린이 효과를 보인다.[54]

진화

조류 메타뉴모바이러스가 1970년대에 보고된 것에 비해, hMPV는 2001년이 되어서야 알려졌다. 베이즈 추론법에 의하면 hMPV는 대략 1800년대에 조류 메타뉴모바이러스에서 분리되었으며 궁극적으로는 119년에서 133년 전에 발생한 것으로 보인다.[55]

각주

  1. ↑ “ICTV Online (10th) Report”. 
  2. ↑ 가 나 van den Hoogen, Bernadette G.; Jong, Jan C. de; Groen, Jan; Kuiken, Thijs; Groot, Ronald de; Fouchier, Ron A.M.; Osterhaus, Albert D.M.E. (2001). “A newly discovered human pneumovirus isolated from young children with respiratory tract disease”. 《Nature Medicine》 7 (6): 719–724. doi:10.1038/89098. PMID 11385510. 
  3. ↑ Amarasinghe, Gaya K.; Bào, Yīmíng; Basler, Christopher F.; Bavari, Sina; Beer, Martin; Bejerman, Nicolás; Blasdell, Kim R.; Bochnowski, Alisa; Briese, Thomas (2017년 4월 7일). “Taxonomy of the order Mononegavirales: update 2017”. 《Archives of Virology》 162 (8): 2493–2504. doi:10.1007/s00705-017-3311-7. PMC 5831667. PMID 28389807. 
  4. ↑ 가 나 다 Williams, John V.; Harris, Paul A.; Tollefson, Sharon J.; Halburnt-Rush, Lisa L.; Pingsterhaus, Joyce M.; Edwards, Kathryn M.; Wright, Peter F.; Crowe, James E. Jr. (2004년 1월 29일). “Human Metapneumovirus and Lower Respiratory Tract Disease in Otherwise Healthy Infants and Children”. 《New England Journal of Medicine》 350 (5): 443–450. doi:10.1056/nejmoa025472. ISSN 0028-4793. PMC 1831873. PMID 14749452. 
  5. ↑ Boivin, Guy; Serres, Gaston De; Côté, Stéphanie; Gilca, Rodica; Abed, Yacine; Rochette, Louis; Bergeron, Michel G.; Déry, Pierre (2003). “Human Metapneumovirus Infections in Hospitalized Children1”. 《Emerging Infectious Diseases》 (미국 영어) 9 (6): 634–640. doi:10.3201/eid0906.030017. PMC 3000156. PMID 12781001. 
  6. ↑ Edwards, Kathryn M.; Zhu, Yuwei; Griffin, Marie R.; Weinberg, Geoffrey A.; Hall, Caroline B.; Szilagyi, Peter G.; Staat, Mary A.; Iwane, Marika; Prill, Mila M. (2013년 2월 13일). “Burden of Human Metapneumovirus Infection in Young Children”. 《New England Journal of Medicine》 (영어) 368 (7): 633–643. doi:10.1056/nejmoa1204630. PMC 3662802. PMID 23406028. 
  7. ↑ Peiris, J.S. Malik; Tang, Wing-Hong; Chan, Kwok-Hung; Khong, Pek-Lan; Guan, Yi; Lau, Yu-Lung; Chiu, Susan S. (2003). “Children with Respiratory Disease Associated with Metapneumovirus in Hong Kong”. 《Emerging Infectious Diseases》 (미국 영어) 9 (6): 628–633. doi:10.3201/eid0906.030009. PMC 3000155. PMID 12781000. 
  8. ↑ Ebihara, Takashi; Endo, Rika; Kikuta, Hideaki; Ishiguro, Nobuhisa; Ishiko, Hiroaki; Hara, Michimaru; Takahashi, Yutaka; Kobayashi, Kunihiko (January 2004). “Human metapneumovirus infection in Japanese children”. 《Journal of Clinical Microbiology》 42 (1): 126–132. doi:10.1128/jcm.42.1.126-132.2004. ISSN 0095-1137. PMC 321731. PMID 14715742. 
  9. ↑ Choi, Eun Hwa; Lee, Hoan Jong; Kim, Sun Jung; Eun, Byung Wook; Kim, Nam Hee; Lee, Jin A.; Lee, Jun Ho; Song, Eun Kyung; Park, So Hee Kim1 Ji Yong (2006년 9월 1일). “The Association of Newly Identified Respiratory Viruses with Lower Respiratory Tract Infections in Korean Children, 2000–2005”. 《Clinical Infectious Diseases》 43 (5): 585–592. doi:10.1086/506350. ISSN 1058-4838. PMID 16886150. 
  10. ↑ Sloots, Theo P.; Mackay, Ian M.; Bialasiewicz, Seweryn; Jacob, Kevin C.; McQueen, Emily; Harnett, Gerald B.; Siebert, David J.; Masters, Brent I.; Young, Paul R. (August 2006). “Human metapneumovirus, Australia, 2001-2004”. 《Emerging Infectious Diseases》 12 (8): 1263–1266. doi:10.3201/eid1708.051239. ISSN 1080-6040. PMC 3291208. PMID 16965711. 
  11. ↑ Madhi, Shabir A.; Ludewick, Herbert; Kuwanda, Locadiah; Niekerk, Nadia van; Cutland, Clare; Klugman, Keith P. (2007). “Seasonality, Incidence, and Repeat Human Metapneumovirus Lower Respiratory Tract Infections in an Area With a High Prevalence of Human Immunodeficiency Virus Type-1 Infection”. 《The Pediatric Infectious Disease Journal》 26 (8): 693–699. doi:10.1097/inf.0b013e3180621192. PMID 17848880. 
  12. ↑ Anderson, Evan J.; Simões, Eric A. F.; Buttery, Jim P.; Dennehy, Penelope H.; Domachowske, Joseph B.; Jensen, Kathryn; Lieberman, Jay M.; Losonsky, Genevieve A.; Yogev, Ram (2012년 9월 1일). “Prevalence and Characteristics of Human Metapneumovirus Infection Among Hospitalized Children at High Risk for Severe Lower Respiratory Tract Infection”. 《Journal of the Pediatric Infectious Diseases Society》 1 (3): 212–222. doi:10.1093/jpids/pis069. ISSN 2048-7193. PMID 26619409. 
  13. ↑ Jain, Seema; Williams, Derek J.; Arnold, Sandra R.; Ampofo, Krow; Bramley, Anna M.; Reed, Carrie; Stockmann, Chris; Anderson, Evan J.; Grijalva, Carlos G. (2015년 2월 25일). “Community-Acquired Pneumonia Requiring Hospitalization among U.S. Children”. 《New England Journal of Medicine》 (영어) 372 (9): 835–845. doi:10.1056/nejmoa1405870. PMC 4697461. PMID 25714161. 
  14. ↑ Aberle, Judith H.; Aberle, Stephan W.; Redlberger-Fritz, Monika; Sandhofer, Michael J.; Popow-Kraupp, Therese (2010). “Human Metapneumovirus Subgroup Changes and Seasonality During Epidemics”. 《The Pediatric Infectious Disease Journal》 29 (11): 1016–8. doi:10.1097/inf.0b013e3181e3331a. PMID 20489673. 
  15. ↑ Williams, John V.; Wang, Chiaoyin K.; Yang, Chin-Fen; Tollefson, Sharon J.; House, Frances S.; Heck, Josh M.; Chu, Marla; Brown, Jennifer B.; Lintao, Linda D. (2006년 2월 1일). “The Role of Human Metapneumovirus in Upper Respiratory Tract Infections in Children: A 20-Year Experience”. 《The Journal of Infectious Diseases》 193 (3): 387–395. doi:10.1086/499274. ISSN 0022-1899. PMC 1586246. PMID 16388486. 
  16. ↑ Ebihara, Takashi; Endo, Rika; Kikuta, Hideaki; Ishiguro, Nobuhisa; Yoshioka, Mikio; Ma, Xiaoming; Kobayashi, Kunihiko (2003년 6월 1일). “Seroprevalence of human metapneumovirus in Japan”. 《Journal of Medical Virology》 (영어) 70 (2): 281–283. doi:10.1002/jmv.10391. ISSN 1096-9071. PMID 12696118. 
  17. ↑ Wolf, Dana G.; Zakay-Rones, Zichria; Fadeela, Ayman; Greenberg, David; Dagan, Ron (2003년 12월 15일). “High Seroprevalence of Human Metapneumovirus among Young Children in Israel”. 《The Journal of Infectious Diseases》 188 (12): 1865–1867. doi:10.1086/380100. ISSN 0022-1899. PMID 14673765. 
  18. ↑ Hamelin, Marie-Eve; Boivin, Guy (February 2005). “Development and validation of an enzyme-linked immunosorbent assay for human metapneumovirus serology based on a recombinant viral protein”. 《Clinical and Diagnostic Laboratory Immunology》 12 (2): 249–253. doi:10.1128/CDLI.12.2.249-253.2005. ISSN 1071-412X. PMC 549303. PMID 15699418. 
  19. ↑ Leung, Jessica; Esper, Frank; Weibel, Carla; Kahn, Jeffrey S. (2005년 3월 1일). “Seroepidemiology of Human Metapneumovirus (hMPV) on the Basis of a Novel Enzyme-Linked Immunosorbent Assay Utilizing hMPV Fusion Protein Expressed in Recombinant Vesicular Stomatitis Virus”. 《Journal of Clinical Microbiology》 (영어) 43 (3): 1213–1219. doi:10.1128/jcm.43.3.1213-1219.2005. ISSN 0095-1137. PMC 1081231. PMID 15750086. 
  20. ↑ 가 나 Pavlin, Julie A.; Hickey, Andrew C.; Ulbrandt, Nancy; Chan, Yee-Peng; Endy, Timothy P.; Boukhvalova, Marina S.; Chunsuttiwat, Supamit; Nisalak, Ananda; Libraty, Daniel H. (2008년 9월 15일). “Human Metapneumovirus Reinfection among Children in Thailand Determined by ELISA Using Purified Soluble Fusion Protein”. 《The Journal of Infectious Diseases》 198 (6): 836–842. doi:10.1086/591186. ISSN 0022-1899. PMC 2648801. PMID 18680407. 
  21. ↑ Dunn, Sarah R.; Ryder, Alex B.; Tollefson, Sharon J.; Xu, Meng; Saville, Benjamin R.; Williams, John V. (2013년 10월 1일). “Seroepidemiologies of Human Metapneumovirus and Respiratory Syncytial Virus in Young Children, Determined with a New Recombinant Fusion Protein Enzyme-Linked Immunosorbent Assay”. 《Clinical and Vaccine Immunology》 (영어) 20 (10): 1654–1656. doi:10.1128/cvi.00750-12. ISSN 1556-6811. PMC 3807191. PMID 23945161. 
  22. ↑ Howard, Leigh M.; Edwards, Kathryn M.; Zhu, Yuwei; Griffin, Marie R.; Weinberg, Geoffrey A.; Szilagyi, Peter G.; Staat, Mary A.; Payne, Daniel C.; Williams, John V. (2017). “Clinical Features of Human Metapneumovirus Infection in Ambulatory Children Aged 5–13 Years”. 《Journal of the Pediatric Infectious Diseases Society》 7 (2): 165–168. doi:10.1093/jpids/pix012. PMC 5954304. PMID 28369564. 
  23. ↑ 가 나 다 Falsey, Ann R.; Erdman, Dean; Anderson, Larry J.; Walsh, Edward E. (2003년 3월 1일). “Human Metapneumovirus Infections in Young and Elderly Adults”. 《The Journal of Infectious Diseases》 187 (5): 785–790. doi:10.1086/367901. ISSN 0022-1899. PMID 12599052. 
  24. ↑ Williams, John V.; Maitre, Nathalie (2016년 7월 28일). “Human metapneumovirus in the preterm neonate: current perspectives”. 《Research and Reports in Neonatology》 (영어) 6: 41–49. doi:10.2147/rrn.s76270. PMC 5120728. PMID 27891060. 
  25. ↑ 가 나 Walsh, Edward E.; Peterson, Derick R.; Falsey, Ann R. (2008년 12월 8일). “Human Metapneumovirus Infections in Adults: Another Piece of the Puzzle”. 《Archives of Internal Medicine》 168 (22): 2489–2496. doi:10.1001/archinte.168.22.2489. ISSN 0003-9926. PMC 2783624. PMID 19064834. 
  26. ↑ 가 나 Widmer, Kyle; Zhu, Yuwei; Williams, John V.; Griffin, Marie R.; Edwards, Kathryn M.; Talbot, H. Keipp (2012년 7월 1일). “Rates of Hospitalizations for Respiratory Syncytial Virus, Human Metapneumovirus, and Influenza Virus in Older Adults”. 《The Journal of Infectious Diseases》 206 (1): 56–62. doi:10.1093/infdis/jis309. ISSN 0022-1899. PMC 3415933. PMID 22529314. 
  27. ↑ Shahda, S.; Carlos, W.g.; Kiel, P.j.; Khan, B.a.; Hage, C.a. (2011년 6월 1일). “The human metapneumovirus: a case series and review of the literature”. 《Transplant Infectious Disease》 (영어) 13 (3): 324–328. doi:10.1111/j.1399-3062.2010.00575.x. ISSN 1399-3062. PMC 3107511. PMID 21631655. 
  28. ↑ Chu, Helen Y.; Renaud, Christian; Ficken, Elle; Thomson, Blythe; Kuypers, Jane; Englund, Janet A. (2014년 12월 1일). “Respiratory Tract Infections Due to Human Metapneumovirus in Immunocompromised Children”. 《Journal of the Pediatric Infectious Diseases Society》 3 (4): 286–293. doi:10.1093/jpids/piu100. ISSN 2048-7193. PMC 4240341. PMID 25419459. 
  29. ↑ Seo, Sachiko; Gooley, Ted A.; Kuypers, Jane M.; Stednick, Zachary; Jerome, Keith R.; Englund, Janet A.; Boeckh, Michael (2016년 7월 15일). “Human Metapneumovirus Infections Following Hematopoietic Cell Transplantation: Factors Associated With Disease Progression”. 《Clinical Infectious Diseases》 63 (2): 178–185. doi:10.1093/cid/ciw284. ISSN 1058-4838. PMC 4928387. PMID 27143659. 
  30. ↑ Shah, Dimpy P.; Shah, Pankil K.; Azzi, Jacques M.; Chaer, Firas El; Chemaly, Roy F. (2016). “Human metapneumovirus infections in hematopoietic cell transplant recipients and hematologic malignancy patients: A systematic review”. 《Cancer Letters》 379 (1): 100–106. doi:10.1016/j.canlet.2016.05.035. PMC 4935561. PMID 27260872. 
  31. ↑ Widmer, Kyle; Griffin, Marie R.; Zhu, Yuwei; Williams, John V.; Talbot, H. Keipp (2014년 5월 1일). “Respiratory syncytial virus- and human metapneumovirus-associated emergency department and hospital burden in adults”. 《Influenza and Other Respiratory Viruses》 (영어) 8 (3): 347–352. doi:10.1111/irv.12234. ISSN 1750-2659. PMC 3984605. PMID 24512531. 
  32. ↑ Williams, John V.; Tollefson, Sharon J.; Heymann, Peter W.; Carper, Holliday T.; Patrie, James; Crowe Jr., James E. (2005). “Human metapneumovirus infection in children hospitalized for wheezing”. 《Journal of Allergy and Clinical Immunology》 115 (6): 1311–1312. doi:10.1016/j.jaci.2005.02.001. PMC 1476700. PMID 15940152. 
  33. ↑ García-García, M.l.; Calvo, C.; Casas, I.; Bracamonte, T.; Rellán, A.; Gozalo, F.; Tenorio, T.; Pérez-Breña, P. (2007년 5월 1일). “Human metapneumovirus bronchiolitis in infancy is an important risk factor for asthma at age 5”. 《Pediatric Pulmonology》 (영어) 42 (5): 458–464. doi:10.1002/ppul.20597. ISSN 1099-0496. PMID 17427899. 
  34. ↑ Khetsuriani, Nino; Kazerouni, N. Neely; Erdman, Dean D.; Lu, Xiaoyan; Redd, Stephen C.; Anderson, Larry J.; Teague, W. Gerald (2007). “Prevalence of viral respiratory tract infections in children with asthma”. 《Journal of Allergy and Clinical Immunology》 119 (2): 314–321. doi:10.1016/j.jaci.2006.08.041. PMID 17140648. 
  35. ↑ Williams, John V.; Crowe, James E.; Enriquez, Rachel; Minton, Patricia; Peebles, R. Stokes; Hamilton, Robert G.; Higgins, Stanley; Griffin, Marie; Hartert, Tina V. (2005년 10월 1일). “Human Metapneumovirus Infection Plays an Etiologic Role in Acute Asthma Exacerbations Requiring Hospitalization in Adults”. 《The Journal of Infectious Diseases》 192 (7): 1149–1153. doi:10.1086/444392. ISSN 0022-1899. PMC 1476781. PMID 16136455. 
  36. ↑ Vicente, Diego; Montes, Milagrosa; Cilla, Gustavo; Pérez-Trallero, Emilio (July 2004). “Human Metapneumovirus and Chronic Obstructive Pulmonary Disease”. 《Emerging Infectious Diseases》 10 (7): 1338–1339. doi:10.3201/eid1007.030633. ISSN 1080-6040. PMC 3323314. PMID 15338546. 
  37. ↑ Martinello, Richard A.; Esper, Frank; Weibel, Carla; Ferguson, David; Landry, Marie L.; Kahn, Jeffrey S. (2006). “Human metapneumovirus and exacerbations of chronic obstructive pulmonary disease”. 《Journal of Infection》 53 (4): 248–254. doi:10.1016/j.jinf.2005.11.010. PMID 16412516. 
  38. ↑ Kan-o, Keiko; Ramirez, Ruben; MacDonald, Martin I.; Rolph, Michael; Rudd, Penny A.; Spann, Kirsten M.; Mahalingam, Suresh; Bardin, Philip G.; Thomas, Belinda J. (2017년 5월 15일). “Human Metapneumovirus Infection in Chronic Obstructive Pulmonary Disease: Impact of Glucocorticosteroids and Interferon”. 《The Journal of Infectious Diseases》 215 (10): 1536–1545. doi:10.1093/infdis/jix167. ISSN 0022-1899. PMID 28379462. 
  39. ↑ Boivin, Guy; Serres, Gaston De; Hamelin, Marie-Eve; Côté, Stéphanie; Argouin, Marco; Tremblay, Geneviève; Maranda-Aubut, Renée; Sauvageau, Chantal; Ouakki, Manale (2007년 5월 1일). “An Outbreak of Severe Respiratory Tract Infection Due to Human Metapneumovirus in a Long-Term Care Facility”. 《Clinical Infectious Diseases》 44 (9): 1152–1158. doi:10.1086/513204. ISSN 1058-4838. PMID 17407031. 
  40. ↑ Louie, Janice K.; Schnurr, David P.; Pan, Chao-Yang; Kiang, David; Carter, Connie; Tougaw, Sandra; Ventura, Jean; Norman, Agnes; Belmusto, Vivian (2007년 9월 1일). “A Summer Outbreak of Human Metapneumovirus Infection in a Long-Term-Care Facility”. 《The Journal of Infectious Diseases》 196 (5): 705–708. doi:10.1086/519846. ISSN 0022-1899. PMID 17674312. 
  41. ↑ Neu, Natalie; Plaskett, Theresa; Hutcheon, Gordon; Murray, Meghan; Southwick, Karen L.; Saiman, Lisa (June 2012). “Epidemiology of Human Metapneumovirus in a Pediatric Long-Term Care Facility”. 《Infection Control & Hospital Epidemiology》 33 (6): 545–550. doi:10.1086/665727. ISSN 0899-823X. PMID 22561708. 
  42. ↑ “Outbreaks of Human Metapneumovirus in Two Skilled Nursing Facilities — West Virginia and Idaho, 2011–2012”. 《www.cdc.gov》 (영어). 2017년 9월 16일에 확인함. 
  43. ↑ Yang, Zifeng; Suzuki, Akira; Watanabe, Oshi; Okamoto, Michiko; Ohmi, Akira; Huang, Wenbo; Nishimura, Hidekazu (2014). “Outbreak of human metapneumovirus infection in a severe motor-and-intellectual disabilities ward in Japan”. 《Japanese Journal of Infectious Diseases》 67 (4): 318–321. doi:10.7883/yoken.67.318. ISSN 1884-2836. PMID 25056083. 
  44. ↑ 김해인 인턴기자 (2020.02.10). “신종 코로나와도 유사한 '메타뉴모 바이러스' 日 노인 5명 사망해”. 《금강일보》. 
  45. ↑ van den Hoogen, Bernadette G.; Bestebroer, Theo M.; Osterhaus, Albert D. M. E.; Fouchier, Ron A. M. (2002년 3월 30일). “Analysis of the genomic sequence of a human metapneumovirus”. 《Virology》 295 (1): 119–132. doi:10.1006/viro.2001.1355. hdl:1765/3864. ISSN 0042-6822. PMID 12033771. 
  46. ↑ Cseke, G.; Maginnis, M. S.; Cox, R. G.; Tollefson, S. J.; Podsiad, A. B.; Wright, D. W.; Dermody, T. S.; Williams, J. V. (2009). “Integrin  v 1 promotes infection by human metapneumovirus”. 《Proceedings of the National Academy of Sciences》 106 (5): 1566–1571. doi:10.1073/pnas.0801433106. PMC 2629439. PMID 19164533. 
  47. ↑ Chang, A.; Masante, C.; Buchholz, U. J.; Dutch, R. E. (2012). “Human Metapneumovirus (HMPV) Binding and Infection Are Mediated by Interactions between the HMPV Fusion Protein and Heparan Sulfate”. 《Journal of Virology》 86 (6): 3230–3243. doi:10.1128/JVI.06706-11. PMC 3302303. PMID 22238303. 
  48. ↑ Cox, R. G.; Livesay, S. B.; Johnson, M.; Ohi, M. D.; Williams, J. V. (2012). “The Human Metapneumovirus Fusion Protein Mediates Entry via an Interaction with RGD-Binding Integrins”. 《Journal of Virology》 86 (22): 12148–12160. doi:10.1128/JVI.01133-12. PMC 3486500. PMID 22933271. 
  49. ↑ Wei, Y.; Zhang, Y.; Cai, H.; Mirza, A. M.; Iorio, R. M.; Peeples, M. E.; Niewiesk, S.; Li, J. (2014). “Roles of the Putative Integrin-Binding Motif of the Human Metapneumovirus Fusion (F) Protein in Cell-Cell Fusion, Viral Infectivity, and Pathogenesis”. 《Journal of Virology》 88 (8): 4338–4352. doi:10.1128/JVI.03491-13. PMC 3993731. PMID 24478423. 
  50. ↑ Schowalter, R. M.; Smith, S. E.; Dutch, R. E. (2006). “Characterization of Human Metapneumovirus F Protein-Promoted Membrane Fusion: Critical Roles for Proteolytic Processing and Low pH”. 《Journal of Virology》 80 (22): 10931–10941. doi:10.1128/JVI.01287-06. PMC 1642150. PMID 16971452. 
  51. ↑ Cox, Reagan G.; Mainou, Bernardo A.; Johnson, Monika; Hastings, Andrew K.; Schuster, Jennifer E.; Dermody, Terence S.; Williams, John V. (2015). “Human Metapneumovirus is Capable of Entering Cells by Fusion with Endosomal Membranes”. 《PLOS Pathogens》 11 (12): e1005303. doi:10.1371/journal.ppat.1005303. PMC 4667933. PMID 26629703. 
  52. ↑ Peiris, JS; Tang, WH; Chan, KH; Khong, PL; Guan, Y; Lau, YL; Chiu, SS (June 2003). “Children with respiratory disease associated with metapneumovirus in Hong Kong”. 《Emerging Infectious Diseases》 9 (6): 628–633. doi:10.3201/eid0906.030009. PMC 3000155. PMID 12781000. 
  53. ↑ Bao X, Liu T, Shan Y, Li K, Garofalo RP, Casola A (May 2008). Baric RS, 편집. “Human Metapneumovirus Glycoprotein G Inhibits Innate Immune Responses”. 《PLOS Pathog.》 4 (5): e1000077. doi:10.1371/journal.ppat.1000077. PMC 2386556. PMID 18516301. 
  54. ↑ Deffrasnes C, Hamelin ME, Boivin G (April 2007). “Human metapneumovirus”. 《Semin Respir Crit Care Med》 28 (2): 213–21. doi:10.1055/s-2007-976493. PMID 17458775. 
  55. ↑ de Graaf M, Osterhaus AD, Fouchier RA, Holmes EC (2008). “Evolutionary dynamics of human and avian metapneumoviruses”. 《J. Gen. Virol.》 89 (Pt 12): 2933–42. doi:10.1099/vir.0.2008/006957-0. PMID 19008378. 

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