Henipavirus

Henipavirus
Colored transmission electron micrograph of a Hendra virion (ca. 300 nm length)
Virus classification
Group: Group V ((-)ssRNA)
Order: Mononegavirales
Family: Paramyxoviridae
Genus: Henipavirus
Type species
Hendra virus
Species

Cedar henipavirus
Ghanaian bat henipavirus
Hendra virus
Mojiang henipavirus
Nipah virus

Henipavirus is a genus of RNA viruses in the family Paramyxoviridae, order Mononegavirales containing five established species. Henipaviruses are naturally harboured by pteropid fruit bats (flying foxes) and microbats of several species.[1] Henipaviruses are characterised by long genomes, a wide host range, and their recent emergence as zoonotic pathogens capable of causing illness and death in domestic animals and humans is cause of concern.[2]

In 2009, RNA sequences of three novel viruses in phylogenetic relationship to known henipaviruses were detected in African straw-colored fruit bats (Eidolon helvum) in Ghana. The finding of these novel henipaviruses outside Australia and Asia indicates that the region of potential endemicity of henipaviruses may be worldwide.[3] These African henipaviruses are slowly being characterised.[4]

Taxonomy

Genus Henipavirus: species and their viruses[5]
Genus Species Virus (Abbreviation)
Henipavirus Cedar henipavirus Cedar virus (CedV)
Ghanaian bat henipavirus Kumasi virus (KV)
Hendra virus* Hendra virus (HeV)
Mojiang henipavirus Mòjiāng virus (MojV)
Nipah virus Nipah virus (NiV)

Table legend: "*" denotes type species.

Virus structure

Structure of henipaviruses
The henipavirus genome (3’ to 5’ orientation) and products of the P gene

Henipaviriona are pleomorphic (variably shaped), ranging in size from 40 to 600 nm in diameter.[6] They possess a lipid membrane overlying a shell of viral matrix protein. At the core is a single helical strand of genomic RNA tightly bound to N (nucleocapsid) protein and associated with the L (large) and P (phosphoprotein) proteins, which provide RNA polymerase activity during replication.

Embedded within the lipid membrane are spikes of F (fusion) protein trimers and G (attachment) protein tetramers. The function of the G protein is to attach the virus to the surface of a host cell via EFNB2, a highly conserved protein present in many mammals.[7][8][9] The structure of the attachment glycoprotein has been determined by X-ray crystallography.[10] The F protein fuses the viral membrane with the host cell membrane, releasing the virion contents into the cell. It also causes infected cells to fuse with neighbouring cells to form large, multinucleated syncytia.

Genome structure

As all mononegaviral genomes, Hendra virus and Nipah virus genomes are non-segmented, single-stranded negative-sense RNA. Both genomes are 18.2 kb in length and contain six genes corresponding to six structural proteins.[11]

In common with other members of the Paramyxoviridae family, the number of nucleotides in the henipavirus genome is a multiple of six, consistent with what is known as the 'rule of six'.[12] Deviation from the rule of six, through mutation or incomplete genome synthesis, leads to inefficient viral replication, probably due to structural constraints imposed by the binding between the RNA and the N protein.

Henipaviruses employ an unusual process called RNA editing to generate multiple proteins from a single gene. The specific process in henipaviruses involves the insertion of extra guanosine residues into the P gene mRNA prior to translation. The number of residues added determines whether the P, V or W proteins are synthesised. The functions of the V and W proteins are unknown, but they may be involved in disrupting host antiviral mechanisms.

Hendra virus

Emergence

Hendra virus (originally called "Equine morbillivirus" was discovered in September 1994 when it caused the deaths of thirteen horses, and a trainer at a training complex in Hendra, a suburb of Brisbane in Queensland, Australia.[13]

The index case, a mare called Drama Series, was housed with 19 other horses after falling ill, and died two days later. Subsequently, all of the horses became ill, with 13 dying. The remaining six animals were subsequently euthanised as a way of preventing relapsing infection and possible further transmission.[14] The trainer, Victory ('Vic') Rail, and the stable foreman, Ray Unwin, were involved in nursing the index case, and both fell ill with an influenza-like illness within one week of the first horse’s death. The stable hand recovered while Rail died of respiratory and renal failure. The source of the virus was most likely frothy nasal discharge from the index case.[15]

A second outbreak occurred in August 1994 (chronologically preceding the first outbreak) in Mackay 1,000 km north of Brisbane resulting in the deaths of two horses and their owner.[16] The owner, Mark Preston, assisted in necropsies of the horses and within three weeks was admitted to hospital suffering from meningitis. Mr Preston recovered, but 14 months later developed neurologic signs and died. This outbreak was diagnosed retrospectively by the presence of Hendra virus in the brain of the patient.[17]

Transmission

Flying foxes have been identified as the reservoir host of Hendra Virus. A seroprevalence of 47% is found in the flying foxes, suggesting an endemic infection of the bat population throughout Australia.[18] Horses become infected with Hendra after exposure to bodily fluid from an infected flying fox. This often happens in the form of urine, feces, or masticated fruit covered in the flying fox's saliva when horses are allowed to graze below roosting sites. The seven human cases have all been infected only after contact with sick horses. As a result, veterinarians are particularly at risk for contracting the disease.

There is no evidence of bat to human, human to human, or human to horse transmission.

Australian outbreaks

Hendra
Mackay
Townsville
Bowen
Hervey Bay
Chinchilla
Rockhampton
Ingham
Cairns
Outbreaks in Queensland
Ballina 
Macksville
Mullumbimby
Lismore
Kempsey
Outbreaks in New South Wales

As of June 2014, a total of fifty outbreaks of Hendra virus have occurred in Australia, all involving infection of horses. As a result of these events, eighty-three horses have died or been euthanised. A further four died or were euthanised as a result of possible hendra infection.

Case fatality rate in humans is 60% and in horses 75%.[19]

Four of these outbreaks have spread to humans as a result of direct contact with infected horses. On 26 July 2011 a dog living on the Mt Alford property was reported to have HeV antibodies, the first time an animal other than a flying fox, horse, or human has tested positive outside an experimental situation.[20]

These events have all been on the east coast of Australia, with the most northern event at Cairns, Queensland and the event furthest south at Kempsey, NSW. Until the event at Chinchilla, Queensland in July 2011, all outbreak sites had been within the distribution of at least two of the four mainland flying foxes (fruit bats); Little red flying fox, (Pteropus scapulatus), black flying fox, (Pteropus alecto), grey-headed flying fox, (Pteropus poliocephalus) and spectacled flying fox, (Pteropus conspicillatus). Chinchilla is considered to be only within the range of little red flying fox and is west of the Great Dividing Range. This is the furthest west the infection has ever been identified in horses.

The timing of incidents indicates a seasonal pattern of outbreaks. Initially this was thought to possibly related to the breeding cycle of the little red flying foxes. These species typically give birth between April and May.[21][22] Subsequently however, the Spectacled flying fox and the Black flying fox have been identified as the species more likely to be involved in infection spillovers.[23]

Timing of outbreaks also appears more likely during the cooler months when it is possible the temperature and humidity are more favourable to the longer term survival of the virus in the environment.[24]

There is no evidence of transmission to humans directly from bats, and, as such it appears that human infection only occurs via an intermediate host, a horse.[25] Despite this in 2014 the NSW Government sanctioned the destruction of flying fox colonies.[26]

Events of June–August 2011

In the years 1994–2010, fourteen events were recorded. Between 20 June 2011 and 28 August 2011, a further seventeen events were identified, during which twenty-one horses died.

It's not clear why there has been a sudden increase in the number of spillover events between June and August 2011. Typically HeV spillover events are more common between May and October. This time is sometimes called "Hendra Season",[94] which is a time when there are large numbers of fruit bats of all species congregated in SE Queensland's valuable winter foraging habitat. The weather (warm and humid) is favourable to the survival of henipavirus in the environment.[95]

It is possible flooding in SE Queensland and Northern NSW in December 2010 and January 2011 may have affected the health of the fruit bats. Urine sampling in flying fox camps indicate that a larger proportion of flying foxes than usual are shedding live virus. Biosecurity Queensland's ongoing surveillance usually shows 7% of the animals are shedding live virus. In June and July nearly 30% animals have been reported to be shedding live virus.[96] Present advice is that these events are not being driven by any mutation in HeV itself.[97]

Other suggestions include that an increase in testing has led to an increase in detection. As the actual mode of transmission between bats and horses has not been determined, it is not clear what, if any, factors can increase the chance of infection in horses.[98]

Following the confirmation of a dog with HeV antibodies, on 27 July 2011, the Queensland and NSW governments will boost research funding into the Hendra virus by $6 million to be spent by 2014–2015. This money will be used for research into ecological drivers of infection in the bats and the mechanism of virus transmission between bats and other species.[99][100] A further 6 million dollars was allocated by the federal government with the funds being split, half for human health investigations and half for animal health and biodiversity research.[101]

Prevention, detection and treatment

Three main approaches are currently followed to reduce the risk to humans.[102]

In November 2012, a vaccine became available for horses. The vaccine is to be used in horses only, since, according to CSIRO veterinary pathologist Dr Deborah Middleton, breaking the transmission cycle from flying foxes to horses prevents it from passing to humans, as well as, "a vaccine for people would take many more years."[103][104]
The vaccine is a subunit vaccine that neutralises Hendra virus and is composed of a soluble version of the G surface antigen on Hendra virus and has been successful in ferret models.[105][106][107]
By December 2014, about 300 000 doses had been administered to more than 100 000 horses. About 3 in 1000 had reported incidents; the majority being localised swelling at the injection site. There had been no reported deaths.[108]
In August 2015, The Australian Pesticides and Veterinary Medicines Authority (APVMA) registered the vaccine. In its statement the Australian government agency released all its data on reported side effects.[109] In January 2016, APVMA approved its use in pregnant mares.[110]
Although the research on the Hendra virus detection is ongoing, a promising result has found using antibody-conjugated magnetic particles and quantum dots.[111][112]
Nipah virus and Hendra virus are closely related paramyxoviruses that emerged from bats during the 1990s to cause deadly outbreaks in humans and domesticated animals. National Institute of Allergy and Infectious Diseases (NIAID)-supported investigators developed vaccines for Nipah and Hendra virus based on the soluble G-glycoproteins of the viruses formulated with adjuvants. Both vaccines have been shown to induce strong neutralizing antibodies in different laboratory animals.
Trials began in 2015 to evaluate a monoclonal antibody to be used as a possible complementary treatment for humans exposed to Hendra virus infected horses.[113]

Pathology

Flying foxes experimentally infected with the Hendra virus develop a viraemia then excrete the virus in their urine, faeces and saliva for approximately one week. Although they excrete active virus during this time there is no other indication of an illness.[114] Symptoms of Hendra virus infection of humans may be respiratory, including hemorrhage and edema of the lungs, or encephalitic, resulting in meningitis. In horses, infection usually causes pulmonary oedema, congestion and / or neurological signs.[115]

Ephrin B2 has been identified as the main receptor for the henipaviruses.[116]

Hendra virus has been classified as a Bio-safety Level 4 Hot Agent.

2016 Brisbane Research Conference

In late May, early June 2016, "Hendra scientists, vets and health officers met in Brisbane to bring together all the research into the bat-borne disease."[117]

Nipah virus

Emergence

Pteropus vampyrus (Large flying fox), one of the natural reservoirs of Nipah virus

Nipah virus was identified in April 1999, when it caused an outbreak of neurological and respiratory disease on pig farms in peninsular Malaysia, resulting in 257 human cases, including 105 human deaths and the culling of one million pigs.[16][118] In Singapore, 11 cases, including one death, occurred in abattoir workers exposed to pigs imported from the affected Malaysian farms. The Nipah virus has been classified by the Centers for Disease Control and Prevention as a Category C agent.[119] The name "Nipah" refers to the place, Kampung Baru Sungai Nipah in Negeri Sembilan State, Malaysia, the source of the human case from which Nipah virus was first isolated.[120][121]

The outbreak was originally mistaken for Japanese encephalitis (JE), however, physicians in the area noted that persons who had been vaccinated against JE were not protected, and the number of cases among adults was unusual[122] Despite the fact that these observations were recorded in the first month of the outbreak, the Ministry of Health failed to react accordingly, and instead launched a nationwide campaign to educate people on the dangers of JE and its vector, Culex mosquitoes.

Symptoms of infection from the Malaysian outbreak were primarily encephalitic in humans and respiratory in pigs. Later outbreaks have caused respiratory illness in humans, increasing the likelihood of human-to-human transmission and indicating the existence of more dangerous strains of the virus.

Based on seroprevalence data and virus isolations, the primary reservoir for Nipah virus was identified as Pteropid fruit bats, including Pteropus vampyrus (Large Flying Fox), and Pteropus hypomelanus (Small flying fox), both of which occur in Malaysia.

The transmission of Nipah virus from flying foxes to pigs is thought to be due to an increasing overlap between bat habitats and piggeries in peninsular Malaysia. At the index farm, fruit orchards were in close proximity to the piggery, allowing the spillage of urine, faeces and partially eaten fruit onto the pigs.[123] Retrospective studies demonstrate that viral spillover into pigs may have been occurring in Malaysia since 1996 without detection.[16] During 1998, viral spread was aided by the transfer of infected pigs to other farms, where new outbreaks occurred.

Evolution

The most likely origin of this virus was in 1947 (95% credible interval: 1888-1988).[124] There are two clades of this virus - one with its origin in 1995 (95% credible interval: 1985-2002) and a second with its origin in 1985 (95% credible interval: 1971-1996). The mutation rate was estimated to be 6.5 × 10−4 substitution/site/year (95% credible interval: 2.3 × 10−4 -1.18 × 10−3) similar to other RNA viruses.

Outbreaks

Locations of henipavirus outbreaks (red stars–Hendra virus; blue stars–Nipah virus) and distribution of henipavirus flying fox reservoirs (red shading–Hendra virus ; blue shading–Nipah virus)

Eight more outbreaks of Nipah virus have occurred since 1998, all within Bangladesh and neighbouring parts of India. The outbreak sites lie within the range of Pteropus species (Pteropus giganteus). As with Hendra virus, the timing of the outbreaks indicates a seasonal effect. Cases occurring in Bangladesh during the winters of 2001, 2003, and 2004, were determined to have been caused by the Nipah virus.[125] In February 2011, a Nipah outbreak began at Hatibandha Upazila in the Lalmonirhat District of northern Bangladesh. To date (7 February 2011), there have been 24 cases and 17 deaths in this outbreak.[126]

Nipah virus has been isolated from Lyle's flying fox (Pteropus lylei) in Cambodia[135] and viral RNA found in urine and saliva from P. lylei and Horsfield's roundleaf bat (Hipposideros larvatus) in Thailand.[136] Infective virus has also been isolated from environmental samples of bat urine and partially eaten fruit in Malaysia.[137] Antibodies to henipaviruses have also been found in fruit bats in Madagascar (Pteropus rufus, Eidolon dupreanum)[138] and Ghana (Eidolon helvum)[139] indicating a wide geographic distribution of the viruses. No infection of humans or other species have been observed in Cambodia, Thailand or Africa thus far.

Pathology

In humans, the infection presents as fever, headache and drowsiness. Cough, abdominal pain, nausea, vomiting, weakness, problems with swallowing and blurred vision are relatively common. About a quarter of the patients have seizures and about 60% become comatose and might need mechanical ventilation. In patients with severe disease, their conscious state may deteriorate and they may develop severe hypertension, fast heart rate, and very high temperature.

Nipah virus is also known to cause relapse encephalitis. In the initial Malaysian outbreak, a patient presented with relapse encephalitis some 53 months after his initial infection. There is no definitive treatment for Nipah encephalitis, apart from supportive measures, such as mechanical ventilation and prevention of secondary infection. Ribavirin, an antiviral drug, was tested in the Malaysian outbreak, and the results were encouraging, though further studies are still needed.

While no vaccine currently exists, a recent (2012) study of a trial vaccine developed using the outer proteins of Hendra virus was shown to induce protection against Nipah in African Green Monkeys.[140]

In animals, especially in pigs, the virus causes a porcine respiratory and neurologic syndrome, locally known as "barking pig syndrome" or "one mile cough."

Ephrin B2 has been identified as the main receptor for the henipaviruses.[116]

Cedar virus

Emergence

Cedar Virus (CedV) was first identified in pteropid urine during work on Hendra virus undertaken in Queensland in 2009.[141]

Although the virus is reported to be very similar to both Hendra and Nipah viruses, it does not cause illness in laboratory animals usually susceptible to paramyxoviruses. Animals were able to mount an effective response and create effective antibodies.[141]

The scientists who identified the virus report:

Hendra and Nipah viruses are 2 highly pathogenic paramyxoviruses that have emerged from bats within the last two decades. Both are capable of causing fatal disease in both humans and many mammal species. Serological and molecular evidence for henipa-like viruses have been reported from numerous locations including Asia and Africa, however, until now no successful isolation of these viruses have been reported. This paper reports the isolation of a novel paramyxovirus, named Cedar virus, from fruit bats in Australia. Full genome sequencing of this virus suggests a close relationship with the henipaviruses. Antibodies to Cedar virus were shown to cross react with, but not cross neutralize Hendra or Nipah virus. Despite this close relationship, when Cedar virus was tested in experimental challenge models in ferrets and guinea pigs, we identified virus replication and generation of neutralizing antibodies, but no clinical disease was observed. As such, this virus provides a useful reference for future reverse genetics experiments to determine the molecular basis of the pathogenicity of the henipaviruses.[141]

Causes of emergence

The emergence of henipaviruses parallels the emergence of other zoonotic viruses in recent decades. SARS coronavirus, Australian bat lyssavirus, Menangle virus and probably Ebola virus and Marburg virus are also harbored by bats and are capable of infecting a variety of other species. The emergence of each of these viruses has been linked to an increase in contact between bats and humans, sometimes involving an intermediate domestic animal host. The increased contact is driven both by human encroachment into the bats’ territory (in the case of Nipah, specifically pigpens in said territory) and by movement of bats towards human populations due to changes in food distribution and loss of habitat.

There is evidence that habitat loss for flying foxes, both in South Asia and Australia (particularly along the east coast) as well as encroachment of human dwellings and agriculture into the remaining habitats, is creating greater overlap of human and flying fox distributions.

See also

References

  1. ↑ Li, Y; Wang, J; Hickey, AC; Zhang, Y; Li, Y; Wu, Y; Zhang, Huajun; et al. (December 2008). "Antibodies to Nipah or Nipah-like viruses in bats, China [letter]". Emerging Infectious Diseases. 14 (12): 1974–6. doi:10.3201/eid1412.080359. PMID 19046545.
  2. ↑ Sawatsky (2008). "Hendra and Nipah Virus". Animal Viruses: Molecular Biology. Caister Academic Press. ISBN 978-1-904455-22-6.
  3. ↑ Drexler JF, Corman VM, Gloza-Rausch F, Seebens A, Annan A (2009). Markotter, Wanda, ed. "Henipavirus RNA in African Bats". PLoS ONE. 4 (7): e6367. Bibcode:2009PLoSO...4.6367D. doi:10.1371/journal.pone.0006367.
  4. ↑ Drexler JF, Corman VM, et al. Bats host major mammalian paramyxoviruses. Nat Commun. 2012 Apr 24;3:796. doi:10.1038/ncomms1796
  5. ↑ Afonso, Claudio L.; Amarasinghe, Gaya K.; Bányai, Krisztián; Bào, Yīmíng; Basler, Christopher F.; Bavari, Sina; Bejerman, Nicolás; Blasdell, Kim R.; Briand, François-Xavier (2016-08-01). "Taxonomy of the order Mononegavirales: update 2016". Archives of Virology. 161 (8): 2351–2360. doi:10.1007/s00705-016-2880-1. PMC 4947412Freely accessible. PMID 27216929.
  6. ↑ Hyatt AD, Zaki SR, Goldsmith CS, Wise TG, Hengstberger SG; Zaki; Goldsmith; Wise; Hengstberger (2001). "Ultrastructure of Hendra virus and Nipah virus within cultured cells and host animals". Microbes and Infection. 3 (4): 297–306. doi:10.1016/S1286-4579(01)01383-1. PMID 11334747.
  7. ↑ Bonaparte, M; Dimitrov, A; Bossart, K (2005). "Ephrin-B2 ligand is a functional receptor for Hendra virus and Nipah virus". Proceedings of the National Academy of Sciences. 102 (30): 10652–7. Bibcode:2005PNAS..10210652B. doi:10.1073/pnas.0504887102. PMC 1169237Freely accessible. PMID 15998730.
  8. ↑ Negrete OA, Levroney EL, Aguilar HC (2005). "EphrinB2 is the entry receptor for Nipah virus, an emergent deadly paramyxovirus". Nature. 436 (7049): 401–5. Bibcode:2005Natur.436..401N. doi:10.1038/nature03838. PMID 16007075.
  9. ↑ Bowden, Thomas A.; Crispin, Max; Jones, E. Yvonne; Stuart, David I. (2010-10-01). "Shared paramyxoviral glycoprotein architecture is adapted for diverse attachment strategies". Biochemical Society Transactions. 38 (5): 1349–1355. doi:10.1042/BST0381349. PMC 3433257Freely accessible. PMID 20863312.
  10. ↑ Bowden, Thomas A.; Crispin, Max; Harvey, David J.; Aricescu, A. Radu; Grimes, Jonathan M.; Jones, E. Yvonne; Stuart, David I. (2008-12-01). "Crystal Structure and Carbohydrate Analysis of Nipah Virus Attachment Glycoprotein: a Template for Antiviral and Vaccine Design". Journal of Virology. 82 (23): 11628–11636. doi:10.1128/JVI.01344-08. PMC 2583688Freely accessible. PMID 18815311.
  11. ↑ Wang L, Harcourt BH, Yu M (2001). "Molecular biology of Hendra and Nipah viruses". Microbes and Infection. 3 (4): 279–87. doi:10.1016/S1286-4579(01)01381-8. PMID 11334745.
  12. ↑ Kolakofsky, D; Pelet, T; Garcin, D; Hausmann, S; Curran, J; Roux, L (February 1998). "Paramyxovirus RNA synthesis and the requirement for hexamer genome length: the rule of six revisited". Journal of Virology. 72 (2): 891–9. PMC 124558Freely accessible. PMID 9444980.
  13. 1 2 Selvey LA, Wells RM, McCormack JG (1995). "Infection of humans and horses by a newly described morbillivirus". Medical Journal of Australia. 162 (12): 642–5. PMID 7603375.
  14. 1 2 3 4 5 "Hendra virus: the initial research". Department of Employment, Economic Development, and Innovation, Queensland Primary Industries and Fisheries. Archived from the original on 13 December 2009. Retrieved 27 January 2014.
  15. ↑ Peacock, Mark (29 October 1995). "Outbreak At Victory Lodge". Retrieved 10 January 2015.
  16. 1 2 3 4 Field, H; Young, P; Yob, JM; Mills, J; Hall, L; MacKenzie, J (2001). "The natural history of Hendra and Nipah viruses". Microbes and Infection. 3 (4): 307–14. doi:10.1016/S1286-4579(01)01384-3. PMID 11334748.
  17. ↑ Walker, Jamie (23 July 2011). "Hendra death toll hits 13 for month". The Australian. Retrieved 24 July 2011.
  18. ↑ Quammen, David. Spillover: Animal Infections and the next Human Pandemic. New York: W.W. Norton, 2012. Print.
  19. ↑ Field, H; de Jong, C; Melville, D; Smith, C; Smith, I; Broos, A; Kung, YH; McLaughlin, A; Zeddeman, A (2011). Fooks, Anthony R, ed. "Hendra virus infection dynamics in Australian fruit bats". PLoS ONE. 6 (12): e28678. Bibcode:2011PLoSO...628678F. doi:10.1371/journal.pone.0028678.
  20. ↑ "Chief vet says dog hendra case 'unprecedented'". 612 ABC Brisbane. 27 July 2011. Retrieved 27 January 2014.
  21. ↑ "Little Red Flying Fox (Pteropus scapulatus)". Wildlife Preservation Society of Queensland. Retrieved 21 May 2014.
  22. ↑ Plowright, RK; Field, H. E.; et al. (2008). "Reproduction and nutritional stress are risk factors for Hendra virus infection in little red flying foxes (Pteropus scapulatus)". Proceedings of the Royal Society B: Biological Sciences. 275 (1636): 861–869. doi:10.1098/rspb.2007.1260. PMC 2596896Freely accessible. PMID 18198149.
  23. ↑ Smith, C; et al. (17 June 2014). "Flying-Fox Species Density - A Spatial Risk Factor for Hendra Virus Infection in Horses in Eastern Australia". PLoS ONE. 20 (22): 477–478. doi:10.1371/journal.pone.0099965.
  24. ↑ Fogarty, R; Halpin, Kim; et al. (2008). "Henipavirus susceptibility to environmental variables". Virus Research. 132 (1–2): 140–144. doi:10.1016/j.virusres.2007.11.010. PMC 3610175Freely accessible. PMID 18166242.
  25. ↑ Selvey L (28 October 1996). "Screening of Bat Carers for Antibodies to Equine Morbillivirus" (PDF). CDI. 20 (22): 477–478.
  26. ↑ Pain, Stephanie (17 October 2015). "The real batman". New Scientist. 228 (3043): 47. doi:10.1016/s0262-4079(15)31425-1.
  27. ↑ Field, HE; Barratt, PC; Hughes, RJ; Shield, J; Sullivan, ND (2000). "A fatal case of Hendra virus infection in a horse in north Queensland: clinical and epidemiological features". Australian Veterinary Journal. 78 (4): 279–80. doi:10.1111/j.1751-0813.2000.tb11758.x. PMID 10840578.
  28. 1 2 3 Hanna, JN; McBride, WJ; Brookes, DL (2006). "Hendra virus infection in a veterinarian". Medical Journal of Australia. 185 (10): 562–4. PMID 17115969.
  29. ↑ "Notifiable Diseases: Hendra Virus" (PDF). Animal Health Surveillance. NSW Department of Primary Industries. 4: 4–5. 2006. Retrieved 6 February 2016.
  30. ↑ ProMED-mail. Hendra virus, human, equine – Australia (Queensland) (03): correction. ProMED-mail 2007; 3 September: 20070903.2896.
  31. ↑ "Queensland vet dies from Hendra virus". Australian Broadcasting Corporation. 21 August 2008.
  32. ↑ Brown, Kimberly S (23 August 2008). "Horses and Human Die in Australia Hendra Outbreak; Government Comes Under Fire". The Horse. Retrieved 26 August 2008.
  33. ↑ Perkins, Nigel (2 December 2008) "Independent review of Hendra virus cases at Redlands and Proserpine in July and August 2008". (the 2008 Perkins Review). dpi.qld.gov.au
  34. ↑ "Vet tests positive to Hendra virus". The Australian. 20 August 2009. Retrieved 20 August 2009.
  35. ↑ Natasha Bita (2 September 2009). "Alister Rodgers dies of Hendra virus after 2 weeks in coma". The Australian. Retrieved 2 September 2009.
  36. ↑ "Horse dies from Hendra virus in Queensland". news.com.au. 20 May 2010. Retrieved 20 May 2010.
  37. ↑ "Biosecurity Queensland". Facebook. 1 July 2011. Retrieved 21 May 2014.
  38. ↑ "Biosecurity Queensland confirms second Hendra case in South East Queensland". Queensland Department of Agriculture, Fisheries and Forestry. 2 July 2011. Archived from the original on 2012-03-13.
  39. ↑ "Another confirmed horse with Hendra virus at Mt Alford". Queensland Department of Agriculture, Fisheries and Forestry. 4 July 2011. Retrieved 5 July 2011.
  40. ↑ "Hendra virus infection confirmed in a dog". Facebook. 25 July 2011. Retrieved 26 July 2011.
  41. ↑ Agius, Kym; Marszalek, Jessica; Berry, Petrina (26 July 2011). "Scientists guessing over Hendra dog". The Sydney Morning Herald. Retrieved 26 July 2011.
  42. ↑ Marszalek, Jessica (31 July 2011). "Dog put down after more Hendra tests". The Sydney Morning Herald. Australian Associated Press. Retrieved 31 July 2011.
  43. ↑ Calligeros, Marissa (29 June 2011). "Eight face Hendra tests after horse's death". Brisbane Times. Australian Associated Press. Retrieved 29 June 2011.
  44. ↑ "Biosecurity Queensland adds Logan result to confirmed Hendra cases". Facebook. 21 July 2011. Retrieved 23 July 2011.
  45. ↑ Miles, Janelle; Robertson, Josh (26 July 2011). "Biosecurity Queensland investigates possible Hendra virus case near Chinchilla". The Courier-Mail. Retrieved 24 July 2011.
  46. ↑ "Confirmed case of hendra virus on NSW North Coast". 1 July 2014. Archived from the original on 2012-11-13. Retrieved 3 July 2011.
  47. ↑ Joyce, Jo (11 October 2011). "Hendra horse owners speak out". ABC North Coast NSW. Retrieved 11 October 2011.
  48. ↑ "Biosecurity Bulletin" (PDF) (Press release). NSW Government Department of Primary Industries. 13 July 2011. Retrieved 16 July 2011.
  49. ↑ "Second case of Hendra virus in NSW near Macksville". dpi.nsw.gov.au. 7 July 2011
  50. ↑ "Second horse dies from Hendra in New South Wales on property near Macksville". Australian Associated Press. 7 July 2011. Retrieved 24 July 2011.
  51. ↑ "Update: Hendra virus infection confirmed at Park Ridge" (Press release). Queensland Department of Agriculture, Fisheries and Forestry. 5 July 2011. Retrieved 5 July 2011.
  52. ↑ Hurst, Daniel (12 July 2011). "Hendra outbreak at LNP candidate's horse riding property". Brisbane Times. Australian Associated Press. Retrieved 12 July 2011.
  53. ↑ "Hendra virus case confirmed in Hervey Bay" (Press release). Queensland Department of Agriculture, Fisheries and Forestry. 16 July 2011. Retrieved 16 July 2011.
  54. ↑ "Fourth NSW horse dies from Hendra virus". Australian Associated Press. 17 July 2014. Retrieved 17 July 2011.
  55. ↑ "Hendra virus case confirmed in Boondall area" (Press release). Queensland Department of Agriculture, Fisheries and Forestry. 16 July 2014. Retrieved 16 July 2011.
  56. ↑ "Hendra virus case confirmed in Chinchilla area". Facebook. 23 July 2011. Retrieved 24 July 2011.
  57. ↑ "Fifth Hendra case confirmed at Mullumbimby" (Press release). NSW Department of Primary Industries. 28 July 2011. Retrieved 28 July 2011.
  58. 1 2 "Biosecurity Bulletin" (PDF) (Press release). NSW Department of Primary Industries. 18 August 2011. Retrieved 18 August 2011.
  59. ↑ "Biosecurity Bulletin" (PDF) (Press release). NSW Department of Primary Industries. 17 August 2011. Retrieved 17 August 2011.
  60. ↑ Stephanie Small (23 August 2011). "Another Hendra outbreak in Queensland". Retrieved 23 August 2011.
  61. ↑ "Biosecurity Bulletin" (PDF) (Press release). NSW Department of Primary Industries. 30 August 2011. Retrieved 30 August 2011.
  62. ↑ "New Hendra Virus Case in Caboolture Area". Facebook. 10 October 2011. Retrieved 11 October 2011.
  63. ↑ "Second Hendra virus case confirmed at Beachmere property October 15, 2011 at 1:42am". Facebook Notes. 15 October 2011. Retrieved 15 October 2011.
  64. ↑ "New Hendra virus case in Townsville area". Facebook. 4 January 2012. Retrieved 5 January 2012.
  65. 1 2 "Two new Hendra virus cases confirmed". Facebook. 29 May 2012. Retrieved 30 June 2012.
  66. 1 2 "Quarantine lifted after Hendra outbreaks". Australian Broadcasting Corporation. 12 July 2012. Retrieved 19 July 2012.
  67. ↑ "Hendra virus quarantines lifted in Ingham and Rockhampton". Queensland Department of Agriculture, Fisheries and Forestry. 12 July 2012. Archived from the original on 2013-05-09.
  68. ↑ Guest, Annie (20 July 2014). "Hendra virus found in Rockhampton". PM. Australian Broadcasting Corporation. Retrieved 19 July 2012.
  69. ↑ "Horses put down after showing Hendra symptoms". Australian Broadcasting Corporation. 27 July 2014. Retrieved 27 July 2012.
  70. ↑ "Two horses euthanased on Rockhampton property". Facebook. 26 July 2012. Retrieved 27 July 2012.
  71. ↑ "New Hendra virus case in Mackay". Facebook. 27 June 2012. Retrieved 30 June 2012.
  72. ↑ "New Hendra virus case in Cairns area". Facebook. 27 July 2012. Retrieved 27 July 2012.
  73. ↑ "Testing underway after latest Hendra outbreak". Australian Broadcasting Corporation. 7 September 2012. Retrieved 7 September 2012.
  74. ↑ "New twist to Port Douglas Hendra death". Horse Zone. Retrieved 16 October 2016.
  75. ↑ "Horse dead after contracting Hendra virus". Australian Broadcasting Corporation. 3 November 2012. Retrieved 3 November 2012.
  76. ↑ "Horses to be tested after Hendra virus outbreak". Australian Broadcasting Corporation. 4 November 2012. Retrieved 4 November 2012.
  77. ↑ "Horse dies from Hendra virus near Mackay". Australian Broadcasting Corporation. 23 January 2013. Retrieved 23 January 2013.
  78. ↑ "New Hendra virus case confirmed in Qld". Australian Broadcasting Corporation. 22 February 2013. Retrieved 23 February 2013.
  79. ↑ Honan, Kim (10 June 2013). "First NSW Hendra horse death in two years". Australian Broadcasting Corporation. Retrieved 10 June 2013.
  80. ↑ Burgess, Sam (4 July 2013). "Second Brisbane Valley property faces Hendra virus lockdown". Australian Broadcasting Corporation. Retrieved 4 July 2013.
  81. ↑ Campbell, Camilla (6 July 2013). "Hendra Outbreak Again in Macksville". NBN News. Retrieved 6 July 2013.
  82. ↑ "New Hendra case confirmed on NSW mid north coast" (Press release). NSW Department of Primary Industries. Retrieved 8 July 2013.
  83. ↑ "Dog infected with Hendra". The Land. NSW Department of Primary Industries. 20 July 2013. Retrieved 20 July 2013.
  84. ↑ "New Hendra virus case confirmed on Gold Coast". Facebook. 5 July 2013. Retrieved 6 July 2013.
  85. ↑ "Third Hendra case confirmed west of Kempsey" (Press release). NSW Department of Primary Industries. 8 July 2013. Retrieved 8 July 2013.
  86. ↑ "Hendra virus claims fourth horse death on NSW mid north coast" (Press release). NSW Department of Primary Industries. 10 July 2013. Retrieved 10 July 2013.
  87. ↑ "Fourth horse dies of Hendra virus at Kempsey on NSW mid north coast". Australian Broadcasting Corporation. 10 July 2013. Retrieved 10 July 2013.
  88. 1 2 Jo Skinner (19 May 2014). "Tests reveal Hendra virus in horse on southern Qld property". Australian Broadcasting Corporation. Retrieved 19 May 2014.
  89. 1 2 "Hendra virus outbreak south of Brisbane" – Australian Broadcasting Corporation – Retrieved 4 June 2014.
  90. 1 2 "Hendra virus confirmed on NSW north coast" – NSW Department of Primary Industries – Retrieved 24 June 2015.
  91. ↑ http://www.gladstoneobserver.com.au/news/biosecurity-still-investigating-hendra-virus/2325550/
  92. ↑ "Hendra virus case confirmed after horse dies in North Queensland". Australian Broadcasting Corporation. 24 July 2015. Retrieved 24 July 2014.
  93. ↑ "Horse dead from Hendra near Lismore in northern New South Wales - By Kim Honan". Australian Broadcasting Corporation. 4 September 2015. Retrieved 5 September 2015.
  94. ↑ Epidemiological methodology of communicating higher contagion periods for better common, non-scientific individual understanding
  95. ↑ Fogarty, R. D.; Halpin, K.; Hyatt, A. D.; Daszak, P.; Mungall, B. A. (2008). "Henipavirus susceptibility to environmental variables". Virus Research. 132 (1–2): 140–144. doi:10.1016/j.virusres.2007.11.010. PMC 3610175Freely accessible. PMID 18166242.
  96. ↑ Tony Moore (28 July 2011). "Nearly a third of bats now carry Hendra: researchers". The Sydney Morning Herald. Retrieved 29 July 2011.
  97. ↑ Janelle Miles; Andrew MacDonald; Koren Helbig (29 July 2011). "Fearon family plead with authorities for stay of execution for Hendra positive dog Dusty". The Courier-Mail. Retrieved 28 July 2011.
  98. ↑ Emma Sykes (4 July 2011). "Hendra virus research continues as more horses contract the disease". 612 ABC Brisbane. Retrieved 5 July 2011.
  99. ↑ "New hunt for Hendra in other species". The Sydney Morning Herald. Australian Associated Press. 27 July 2011. Retrieved 27 July 2011.
  100. ↑ Guest, Annie (27 July 2011). "Urgent funds for hendra research". The World Today. Australian Broadcasting Corporation. Retrieved 27 July 2011.
  101. ↑ "Gillard Government helping in response to Hendra". 29 July 2011. Archived from the original on 2012-02-27.
  102. ↑ "Opinion: combating the deadly Hendra virus". CSIRO. 13 May 2011. Archived from the original on 2012-11-24.
  103. ↑ "Equine Henda Virus Vaccine Launched in Australia". The Horse. 1 November 2012. Retrieved 7 November 2012.
  104. ↑ Taylor, John; Guest, Annie (1 November 2012). "Breakthrough Hendra virus vaccine released for horses". Australian Broadcasting Corporation. Retrieved 1 November 2012.
  105. ↑ Pallister, J; Middleton, D; Wang, LF (2011). "A recombinant Hendra virus G glycoprotein-based subunit vaccine protects ferrets from lethal Hendra virus challenge". Vaccine. 29 (24): 5623–30. doi:10.1016/j.vaccine.2011.06.015. PMC 3153950Freely accessible. PMID 21689706.
  106. ↑ Fraser, Kelmeny (24 July 2011). "Hendra virus scientists push for vaccine to be fast-tracked". The Sunday Mail. Queensland. Retrieved 28 July 2011.
  107. ↑ "Hendra vaccine could be ready in 2012". Australian Associated Press. 17 May 2011. Retrieved 28 July 2011.
  108. ↑ "Autopsy carried out on exhumed horse to determine if Hendra vaccine caused its death - By Marty McCarthy". Australian Broadcasting Corporation 30 December 2014. Retrieved 30 December 2014.
  109. ↑ "Chemical regulator registers Hendra vaccine, releases data on reported side effects - By Marty McCarthy". Australian Broadcasting Corporation 5 August 2015. Retrieved 19 August 2015.
  110. ↑ "Hendra vaccine approved for use in pregnant mares - By Kim Honan". Australian Broadcasting Corporation. 29 January 2016. Retrieved 29 January 2016.
  111. ↑ Huw Morgan (20 September 2012). "A 'quantum' step towards on-the-spot Hendra virus detection". news@CSIRO. Retrieved 20 September 2012.
  112. ↑ Lisi F, Falcaro P, Buso B, Hill AJ, Barr JA, Crameri G, Nguyen T-L, Wang L-F, Mulvaney P; Falcaro; Buso; Hill; Barr; Crameri; Nguyen; Wang; Mulvaney (2012). "Rapid Detection of Hendra Virus Using Magnetic Particles and Quantum Dots". Advanced Healthcare Materials. 1 (5): 631–634. doi:10.1002/adhm.201200072. PMID 23184798.
  113. ↑ - By Robin McConchie. "Hendra trials for humans about treatment not prevention". Australian Broadcasting Corporation. Retrieved 2 April 2015.
  114. ↑ Edmondston, Jo; Field, Hume (2009). "Research update: Hendra Virus" (PDF). Australian Biosecurity CRC for Emerging Infectious Disease. Retrieved 17 July 2011.
  115. ↑ Middleton, D. "1Initial experimental characterisation of HeV (Redland Bay 2008) infection in horses" (PDF). Retrieved 17 July 2011.
  116. 1 2 Lee B, Ataman ZA; Ataman (2011). "Modes of paramyxovirus fusion: a Henipavirus perspective". Trends in Microbiology. 19 (8): 389–399. doi:10.1016/j.tim.2011.03.005. PMC 3264399Freely accessible. PMID 21511478.
  117. ↑ "Black and spectacled flying foxes are main Hendra virus carriers, research shows - By Robin McConchie". Australian Broadcasting Corporation. 3 June 2016. Retrieved 4 June 2016.
  118. ↑ Centers for Disease Control and Prevention (CDC) (30 April 1999). "Update: outbreak of Nipah virus—Malaysia and Singapore, 1999". Morbidity and Mortality Weekly Report. United States Centers for Disease Control and Prevention. 48 (16): 335–7. PMID 10366143.
  119. ↑ Bioterrorism Agents/Diseases. bt.cdc.gov
  120. ↑ Siva SR, Chong HT, Tan CT (2009). "Ten year clinical and serological outcomes of Nipah virus infection" (PDF). Neurology Asia. 14: 53–58.
  121. ↑ "Spillover — Zika, Ebola & Beyond". pbs.org. PBS. 3 August 2016. Retrieved 4 August 2016.
  122. ↑ "Dobbs and the viral encephalitis outbreak".. Archived thread from the Malaysian Doctors Only BBS.
  123. ↑ Chua KB, Chua BH, Wang CW; Chua; Wang (2002). "Anthropogenic deforestation, El Niño and the emergence of Nipah virus in Malaysia". The Malaysian Journal of Pathology. 24 (1): 15–21. PMID 16329551.
  124. ↑ Lo Presti A, Cella E, Giovanetti M, Lai A, Angeletti S, Zehender G, Ciccozzi M (2015). "Origin and evolution of Nipah virus". J Med Virol. 88 (3): 380–388. doi:10.1002/jmv.24345. PMID 26252523.
  125. ↑ Chadha MS, Comer JA, Lowe L, Rota PA, Rollin PE, Bellini WJ, Ksiazek TG, Mishra A; Comer; Lowe; Rota; Rollin; Bellini; Ksiazek; Mishra (February 2006). "Nipah virus-associated encephalitis outbreak, Siliguri, India". Emerging Infectious Diseases. 12 (2): 235–40. doi:10.3201/eid1202.051247. PMC 3373078Freely accessible. PMID 16494748.
  126. ↑ "Nipah Outbreak at Lalmonirhat". Bangladesh: Institute of Epidemiology, Disease Control and Research. 7 February 2011. Retrieved 21 May 2014.
  127. ↑ Chadha MS, Comer JA, Lowe L (2006). "Nipah virus-associated encephalitis outbreak, Siliguri, India". Emerging Infectious Diseases. 12 (2): 235–40. doi:10.3201/eid1202.051247. PMC 3373078Freely accessible. PMID 16494748.
  128. 1 2 Hsu VP, Hossain MJ, Parashar UD (2004). "Nipah virus encephalitis reemergence, Bangladesh". Emerging Infectious Diseases. 10 (12): 2082–7. doi:10.3201/eid1012.040701. PMC 3323384Freely accessible. PMID 15663842.
  129. ↑ ICDDR,B (2005). "Nipah virus outbreak from date palm juice". Health and Science Bulletin. 3 (4): 1–5. Archived from the original on 10 December 2006.
  130. ↑ ICDDR,B (2007). "Person-to-person transmission of Nipah infection in Bangladesh". Health and Science Bulletin. 5 (4): 1–6. Archived from the original on 3 February 2009.
  131. ↑ ICDDR,B (2008). "Outbreaks of Nipah virus in Rajbari and Manikgonj". Health and Science Bulletin. 6 (1): 12–3. Archived from the original on 3 February 2009.
  132. ↑ ICDDR,B (2010). "Nipah outbreak in Faridpur District, Bangladesh, 2010". Health and Science Bulletin. 8 (2): 6–11. Archived from the original on 28 September 2011.
  133. ↑ "Arguments in Bahodderhat murder case begin". The Daily Star. 18 March 2008. Retrieved 21 May 2014.
  134. ↑ তাহেরকে ফাঁসি দেওয়ার সিদ্ধান্ত নেন জিয়া. prothom-alo.com. 4 February 2011
  135. ↑ Reynes JM, Counor D, Ong S (2005). "Nipah virus in Lyle's flying foxes, Cambodia". Emerging Infectious Diseases. 11 (7): 1042–7. doi:10.3201/eid1107.041350. PMC 3371782Freely accessible. PMID 16022778.
  136. ↑ Wacharapluesadee S, Lumlertdacha B, Boongird K (2005). "Bat Nipah virus, Thailand". Emerging Infectious Diseases. 11 (12): 1949–51. doi:10.3201/eid1112.050613. PMC 3367639Freely accessible. PMID 16485487.
  137. ↑ Chua KB, Koh CL, Hooi PS (2002). "Isolation of Nipah virus from Malaysian Island flying-foxes". Microbes and Infection. 4 (2): 145–51. doi:10.1016/S1286-4579(01)01522-2. PMID 11880045.
  138. ↑ Lehlé C, Razafitrimo G, Razainirina J (2007). "Henipavirus and Tioman virus antibodies in pteropodid bats, Madagascar". Emerging Infectious Diseases. 13 (1): 159–61. doi:10.3201/eid1301.060791. PMC 2725826Freely accessible. PMID 17370536.
  139. ↑ Hayman D, Suu-Ire R, Breed A; et al. (2008). Montgomery, Joel Mark, ed. "Evidence of henipavirus infection in West African fruit bats". PLoS ONE. 3 (7): 2739. Bibcode:2008PLoSO...3.2739H. doi:10.1371/journal.pone.0002739. PMC 2453319Freely accessible. PMID 18648649.
  140. ↑ Bossart K. N.; Rockx B.; Feldmann F.; Brining D.; Scott D.; LaCasse R.; Geisbert J. B.; Feng Y.-R.; Chan Y.-P. (2012). "A Hendra Virus G Glycoprotein Subunit Vaccine Protects African Green Monkeys from Nipah Virus Challenge". Science Translational Medicine. 4 (146): 146ra107. doi:10.1126/scitranslmed.3004241.
  141. 1 2 3 Marsh, Glenn A.; de Jong, Carol; Barr, Jennifer A.; Tachedjian, Mary; Smith, Craig; Middleton, Deborah; Yu, Meng; Todd, Shawn; Foord, Adam J.; Haring, Volker; Payne, Jean; Robinson, Rachel; Broz, Ivano; Crameri, Gary; Field, Hume E.; Wang, Lin-Fa (2 August 2012). "Cedar Virus: A Novel Henipavirus Isolated from Australian Bats". PLOS Pathogens. 8 (8): e1002836. doi:10.1371/journal.ppat.1002836. PMC 3410871Freely accessible. PMID 22879820.
Wikimedia Commons has media related to Henipavirus.
This article is issued from Wikipedia - version of the 12/1/2016. The text is available under the Creative Commons Attribution/Share Alike but additional terms may apply for the media files.