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{{short description|Occurs when a reservoir population causes an epidemic in a novel host population}}
{{short description|Occurs when a reservoir population causes an epidemic in a novel host population}}
'''Spillover infection''', also known as '''pathogen spillover''' and '''spillover event''', occurs when a [[Natural reservoir|reservoir]] population with a high [[pathogen]] prevalence comes into contact with a novel host population. The pathogen is transmitted from the reservoir population and may or may not be transmitted within the host population. Due to [[climate change]] and land use expansion, the risk of viral spillover is predicted to significantly increase.<ref name="Carlson">{{cite journal |last1=Carlson |first1=Colin J. |last2=Albery |first2=Gregory F. |last3=Merow |first3=Cory |last4=Trisos |first4=Christopher H. |last5=Zipfel |first5=Casey M. |last6=Eskew |first6=Evan A. |last7=Olival |first7=Kevin J. |last8=Ross |first8=Noam |last9=Bansal |first9=Shweta |title=Climate change increases cross-species viral transmission risk |journal=Nature |date=28 April 2022 |volume=607 |issue=7919 |pages=555–562 |doi=10.1038/s41586-022-04788-w|pmid=35483403 |bibcode=2022Natur.607..555C |s2cid=248430532 |language=en |issn=1476-4687|doi-access=free }}</ref><ref>{{cite journal | last1 = Power | first1 = AG | last2 = Mitchell | first2 = CE | date = Nov 2004 | title = Pathogen spillover in disease epidemics | journal = The American Naturalist| volume = 164 | issue = Suppl 5| pages = S79–89 | doi = 10.1086/424610 | pmid = 15540144 | s2cid = 16762851 }}</ref>
'''Spillover infection''', also known as '''pathogen spillover''' and '''spillover event''', occurs when a [[Natural reservoir|reservoir]] population with a high [[pathogen]] prevalence comes into contact with a novel host population. The pathogen is transmitted from the reservoir population and may or may not be transmitted within the host population.<ref name=":2">{{cite journal |last1=Woolhouse |first1=Mark |last2=Scott |first2=Fiona |last3=Hudson |first3=Zoe |last4=Howey |first4=Richard |last5=Chase-Topping |first5=Margo |year=2012 |title=Human viruses: Discovery and emergence |journal=Philosophical Transactions of the Royal Society B: Biological Sciences |volume=367 |issue=1604 |pages=2864–2871 |doi=10.1098/rstb.2011.0354 |pmc=3427559 |pmid=22966141}}</ref> Due to [[climate change]] and land use expansion, the risk of viral spillover is predicted to significantly increase.<ref name="Wolfe">{{cite journal |last1=Wolfe |first1=Nathan D. |last2=Dunavan |first2=Claire Panosian |last3=Diamond |first3=Jared |date=May 2007 |title=Origins of major human infectious diseases |journal=Nature |language=en |volume=447 |issue=7142 |pages=279–283 |bibcode=2007Natur.447..279W |doi=10.1038/nature05775 |issn=1476-4687 |pmc=7095142 |pmid=17507975}}</ref><ref>''Ebola.'' (2014). National Center for Emerging and Zoonotic Infectious Diseases, Division of High-Consequence Pathogens and Pathology, Department of Health & Human Services, CDC.</ref>


== Spillover zoonoses ==
== Spillover zoonoses ==
[[File:Fruit Bat (flying fox) (35908751483).jpg|thumb|The [[fruit bat]] is believed to be the zoonotic agent responsible for the spillover of the [[Ebola virus]].]]
[[File:Fruit Bat (flying fox) (35908751483).jpg|thumb|The [[fruit bat]] is believed to be the zoonotic agent responsible for the spillover of the [[Ebola virus]].]]


Spillover is a common event; in fact, more than two-thirds of human viruses are [[zoonosis|zoonotic]].<ref>{{cite journal | doi=10.1098/rstb.2011.0354 | title=Human viruses: Discovery and emergence | year=2012 | last1=Woolhouse | first1=Mark | last2=Scott | first2=Fiona | last3=Hudson | first3=Zoe | last4=Howey | first4=Richard | last5=Chase-Topping | first5=Margo | journal=Philosophical Transactions of the Royal Society B: Biological Sciences | volume=367 | issue=1604 | pages=2864–2871 | pmid=22966141 | pmc=3427559 }}</ref><ref name="Wolfe">{{cite journal |last1=Wolfe |first1=Nathan D. |last2=Dunavan |first2=Claire Panosian |last3=Diamond |first3=Jared |title=Origins of major human infectious diseases |journal=Nature |date=May 2007 |volume=447 |issue=7142 |pages=279–283 |doi=10.1038/nature05775 |pmid=17507975 |pmc=7095142 |bibcode=2007Natur.447..279W |language=en |issn=1476-4687}}</ref> Most spillover events result in self-limited cases with no further human to human transmission, as occurs, for example, with rabies, anthrax, [[histoplasmosis]] or hidatidosis. Other zoonotic pathogens are able to be transmitted by humans to produce secondary cases and even to establish limited chains of transmission. Some examples are the [[Ebola]] and Marburg filoviruses, the [[MERS]] and [[SARS]] coronaviruses and some avian flu viruses. Finally, some spillover events can result in the final adaptation of the microbe to humans, who can become a new stable reservoir, as occurred with the [[HIV virus]] resulting in the [[Epidemiology_of_HIV/AIDS|AIDS epidemic]] and with [[SARS-CoV-2]] resulting in the [[COVID-19 pandemic]].<ref name="Wolfe"/>
Spillover is a common event; in fact, more than two-thirds of human viruses are [[Zoonosis|zoonotic]].<ref name=":0">{{cite journal |last1=Graystock |first1=P |last2=Yates |first2=K |last3=Evison |first3=SEF |last4=Darvill |first4=B |last5=Goulson |first5=D |last6=Hughes |first6=WOH |year=2013 |title=The Trojan hives: pollinator pathogens, imported and distributed in bumblebee colonies |journal=Journal of Applied Ecology |volume=50 |issue=5 |pages=1207–15 |doi=10.1111/1365-2664.12134 |s2cid=3937352}}</ref><ref name=":1">{{Cite journal |last1=Sachman-Ruiz |first1=Bernardo |last2=Narváez-Padilla |first2=Verónica |last3=Reynaud |first3=Enrique |date=2015-03-10 |title=Commercial Bombus impatiens as reservoirs of emerging infectious diseases in central México |journal=Biological Invasions |volume=17 |issue=7 |pages=2043–53 |doi=10.1007/s10530-015-0859-6 |issn=1387-3547 |doi-access=free}}</ref> Most spillover events result in self-limited cases with no further human-to-human transmission, as occurs, for example, with rabies, anthrax, [[histoplasmosis]] or hydatidosis. Other zoonotic pathogens are able to be transmitted by humans to produce secondary cases and even to establish limited chains of transmission. Some examples are the [[Ebola]] and Marburg filoviruses, the [[MERS]] and [[SARS]] coronaviruses and some avian flu viruses. Finally, some spillover events can result in the final adaptation of the microbe to humans, who can become a new stable reservoir, as occurred with the [[HIV virus]] resulting in the [[Epidemiology of HIV/AIDS|AIDS epidemic]] and with [[SARS-CoV-2]] resulting in the [[COVID-19 pandemic]].<ref name=":1" />


If the history of mutual adaptation is long enough, permanent host-microbe associations can be established resulting in co-evolution, and even permanent integration of the microbe genome with the human genome, as it is the case of endogenous viruses. The closer the two target host species are in phylogenetic terms, the easier it is for microbes to overcome the biological barrier to produce successful spillovers. For this reason, other [[mammals]] are the main source of zoonotic agents for humans. For example, in the case of the [[Ebola]] virus, fruit bats are the hypothesized zoonotic agent.<ref>''Ebola.'' (2014). National Center for Emerging and Zoonotic Infectious Diseases, Division of High-Consequence Pathogens and Pathology, Department of Health & Human Services, CDC.</ref>
If the history of mutual adaptation is long enough, permanent host-microbe associations can be established resulting in co-evolution, and even permanent integration of the microbe genome with the human genome, as is the case of endogenous viruses.<ref>{{Cite journal |last1=Durrer |first1=Stephan |last2=Schmid-Hempel |first2=Paul |date=1994-12-22 |title=Shared Use of Flowers Leads to Horizontal Pathogen Transmission |journal=Proceedings of the Royal Society of London B: Biological Sciences |volume=258 |issue=1353 |pages=299–302 |bibcode=1994RSPSB.258..299D |doi=10.1098/rspb.1994.0176 |issn=0962-8452 |s2cid=84926310}}</ref> The closer the two target host species are in phylogenetic terms, the easier it is for microbes to overcome the biological barrier to produce successful spillovers.<ref name=":2" /> For this reason, other [[mammals]] are the main source of zoonotic agents for humans. For example, in the case of the [[Ebola]] virus, fruit bats are the hypothesized zoonotic agent.<ref>{{Cite journal |last1=Graystock |first1=Peter |last2=Goulson |first2=Dave |last3=Hughes |first3=William O. H. |date=2015-08-22 |title=Parasites in bloom: flowers aid dispersal and transmission of pollinator parasites within and between bee species |journal=Proceedings of the Royal Society B: Biological Sciences |volume=282 |issue=1813 |pages=20151371 |doi=10.1098/rspb.2015.1371 |issn=0962-8452 |pmc=4632632 |pmid=26246556}}</ref>


During the late 20th century zoonotic spillover increased as the [[environmental impact of agriculture]] promoted increased land use and [[deforestation]], changing wildlife [[habitat]]. As species shift their geographic range in response to [[climate change]], the risk of zoonotic spillover is predicted to substantially increase, particularly in tropical regions that are experiencing rapid warming.<ref name="Carlson"/> As forested areas of land are cleared for human use, there is increased proximity and interaction between wild animals and humans thereby increasing the potential for exposure.<ref>{{cite journal | doi=10.1038/srep41613 | title=The nexus between forest fragmentation in Africa and Ebola virus disease outbreaks | year=2017 | last1=Rulli | first1=Maria Cristina | last2=Santini | first2=Monia | last3=Hayman | first3=David T. S. | last4=d'Odorico | first4=Paolo | journal=Scientific Reports | volume=7 | page=41613 | pmid=28195145 | pmc=5307336 | bibcode=2017NatSR...741613R }}</ref>
During the late 20th century, zoonotic spillover increased as the [[environmental impact of agriculture]] promoted increased land use and [[deforestation]], changing wildlife [[habitat]]. As species shift their geographic range in response to [[climate change]], the risk of zoonotic spillover is predicted to substantially increase, particularly in tropical regions that are experiencing rapid warming.<ref>{{cite journal |last1=Otterstatter |first1=MC |last2=Thomson |first2=JD |year=2008 |title=Does Pathogen Spillover from Commercially Reared Bumble Bees Threaten Wild Pollinators? |journal=PLOS ONE |volume=3 |issue=7 |page=e2771 |bibcode=2008PLoSO...3.2771O |doi=10.1371/journal.pone.0002771 |pmc=2464710 |pmid=18648661 |doi-access=free}}</ref> As forested areas of land are cleared for human use, there is increased proximity and interaction between wild animals and humans thereby increasing the potential for exposure.<ref>{{Cite journal |last1=Graystock |first1=Peter |last2=Goulson |first2=Dave |last3=Hughes |first3=William O.H. |year=2014 |title=The relationship between managed bees and the prevalence of parasites in bumblebees |journal=PeerJ |volume=2 |pages=e522 |doi=10.7717/peerj.522 |pmc=4137657 |pmid=25165632 |doi-access=free}}</ref>


== Intraspecies spillover ==
== Intraspecies spillover ==
[[File:Bumblebee closeup.jpg|thumb|The bumblebee is a potential reservoir for several pollinator parasites.|217x217px]]
[[File:Bumblebee closeup.jpg|thumb|The bumblebee is a potential reservoir for several pollinator parasites.|217x217px]]
Commercially bred [[bumblebees]] used to pollinate greenhouses can be reservoirs for several pollinator parasites including the protozoans [[Crithidia|''Crithidia bombi'']], and ''[[Apicystis bombi]]'',<ref name=":0">{{cite journal | last1 = Graystock | first1 = P | last2 = Yates | first2 = K | last3 = Evison | first3 = SEF | last4 = Darvill | first4 = B | last5 = Goulson | first5 = D | last6 = Hughes | first6 = WOH | year = 2013| title = The Trojan hives: pollinator pathogens, imported and distributed in bumblebee colonies | doi = 10.1111/1365-2664.12134 | journal = Journal of Applied Ecology | volume = 50 | issue = 5| pages = 1207–15 | s2cid = 3937352 }}</ref> the microsporidians ''[[Nosema bombi]]'' and ''[[Nosema ceranae]]'',<ref name=":0" /><ref name=":1">{{Cite journal|title = Commercial Bombus impatiens as reservoirs of emerging infectious diseases in central México|journal = Biological Invasions|date = 2015-03-10|issn = 1387-3547|pages = 2043–53|volume = 17|issue = 7|doi = 10.1007/s10530-015-0859-6|first1 = Bernardo|last1 = Sachman-Ruiz|first2 = Verónica|last2 = Narváez-Padilla|first3 = Enrique|last3 = Reynaud|doi-access = free}}</ref> plus viruses such as [[Deformed wing virus]] and the tracheal mites ''[[Locustacarus buchneri]]''.<ref name=":1" /> Commercial bees that escape the glasshouse environment may then infect wild bee populations. Infection may be via direct interactions between managed and wild bees or via shared flower use and contamination.<ref>{{Cite journal|title = Shared Use of Flowers Leads to Horizontal Pathogen Transmission|journal = Proceedings of the Royal Society of London B: Biological Sciences|date = 1994-12-22|issn = 0962-8452|pages = 299–302|volume = 258|issue = 1353|doi = 10.1098/rspb.1994.0176|first1 = Stephan|last1 = Durrer|first2 = Paul|last2 = Schmid-Hempel|bibcode = 1994RSPSB.258..299D|s2cid = 84926310}}</ref><ref>{{Cite journal|title = Parasites in bloom: flowers aid dispersal and transmission of pollinator parasites within and between bee species|journal = Proceedings of the Royal Society B: Biological Sciences|date = 2015-08-22|issn = 0962-8452|pmc = 4632632|pmid = 26246556|pages = 20151371|volume = 282|issue = 1813|doi = 10.1098/rspb.2015.1371|first1 = Peter|last1 = Graystock|first2 = Dave|last2 = Goulson|first3 = William O. H.|last3 = Hughes}}</ref> One study found that half of all wild bees found near greenhouses were infected with ''C. bombi''. Rates and incidence of infection decline dramatically the further away from the greenhouses the wild bees are located.<ref>{{cite journal | last1 = Otterstatter | first1 = MC | last2 = Thomson | first2 = JD | year = 2008| title = Does Pathogen Spillover from Commercially Reared Bumble Bees Threaten Wild Pollinators? | journal = PLOS ONE | volume = 3 | issue = 7| page = e2771 | doi=10.1371/journal.pone.0002771 | pmid=18648661 | pmc=2464710| bibcode = 2008PLoSO...3.2771O | doi-access = free }}</ref><ref>{{Cite journal|title = The relationship between managed bees and the prevalence of parasites in bumblebees|journal = PeerJ|pmc = 4137657|pmid = 25165632|volume = 2|doi = 10.7717/peerj.522|first1 = Peter|last1 = Graystock|first2 = Dave|last2 = Goulson|first3 = William O.H.|last3 = Hughes|pages=e522|year=2014 | doi-access=free }}</ref> Instances of spillover between bumblebees are well documented across the world but particularly in Japan, North America and the United Kingdom.<ref>{{Cite journal|title = Do managed bees drive parasite spread and emergence in wild bees?|journal = International Journal for Parasitology: Parasites and Wildlife|doi = 10.1016/j.ijppaw.2015.10.001|pmid = 28560161|pmc = 5439461|first1 = Peter|last1 = Graystock|first2 = Edward J.|last2 = Blane|first3 = Quinn S.|last3 = McFrederick|first4 = Dave|last4 = Goulson|first5 = William O. H.|last5 = Hughes|volume=5|issue = 1|pages=64–75|year=2016}}</ref><ref>{{cite news |last=Carrington |first=Damian |newspaper=The Guardian|url=https://www.theguardian.com/environment/2013/jul/18/bumblebees-honeybees-parasite-infection|title=Imported bumblebees pose risk to UK's wild and honeybee population – study |date=18 July 2013 }}</ref>
Commercially bred [[bumblebees]] used to pollinate greenhouses can be reservoirs for several pollinator parasites including the protozoans [[Crithidia|''Crithidia bombi'']], and ''[[Apicystis bombi]]'',<ref name=":3" /> the microsporidians ''[[Nosema bombi]]'' and ''[[Nosema ceranae]]'',<ref name=":3" /><ref name=":4" /> plus viruses such as [[Deformed wing virus]] and the tracheal mites ''[[Locustacarus buchneri]]''.<ref name=":4" /> Commercial bees that escape the greenhouse environment may then infect wild bee populations. Infection may be via direct interactions between managed and wild bees or via shared flower use and contamination.<ref name=":5" /><ref>{{Cite journal |last=Graystock |first=Peter |last2=Goulson |first2=Dave |last3=Hughes |first3=William O. H. |date=2015-08-22 |title=Parasites in bloom: flowers aid dispersal and transmission of pollinator parasites within and between bee species |url=https://royalsocietypublishing.org/doi/10.1098/rspb.2015.1371 |journal=Proceedings of the Royal Society B: Biological Sciences |language=en |volume=282 |issue=1813 |pages=20151371 |doi=10.1098/rspb.2015.1371 |issn=0962-8452}}</ref> One study found that half of all wild bees found near greenhouses were infected with ''C. bombi''. Rates and incidence of infection decline dramatically the further away from the greenhouses where the wild bees are located.<ref>{{Cite journal |last=Otterstatter |first=Michael C. |last2=Thomson |first2=James D. |date=2008-07-23 |editor-last=Adler |editor-first=Frederick R. |title=Does Pathogen Spillover from Commercially Reared Bumble Bees Threaten Wild Pollinators? |url=https://dx.plos.org/10.1371/journal.pone.0002771 |journal=PLoS ONE |language=en |volume=3 |issue=7 |pages=e2771 |doi=10.1371/journal.pone.0002771 |issn=1932-6203}}</ref><ref>{{Cite journal |last=Graystock |first=Peter |last2=Goulson |first2=Dave |last3=Hughes |first3=William O.H. |date=2014-08-12 |title=The relationship between managed bees and the prevalence of parasites in bumblebees |url=https://peerj.com/articles/522 |journal=PeerJ |language=en |volume=2 |pages=e522 |doi=10.7717/peerj.522 |issn=2167-8359}}</ref> Instances of spillover between bumblebees are well documented across the world, particularly in Japan, North America, and the United Kingdom.<ref>{{Cite journal |last=Graystock |first=Peter |last2=Blane |first2=Edward J. |last3=McFrederick |first3=Quinn S. |last4=Goulson |first4=Dave |last5=Hughes |first5=William O.H. |date=2016-04 |title=Do managed bees drive parasite spread and emergence in wild bees? |url=https://linkinghub.elsevier.com/retrieve/pii/S2213224415300158 |journal=International Journal for Parasitology: Parasites and Wildlife |language=en |volume=5 |issue=1 |pages=64–75 |doi=10.1016/j.ijppaw.2015.10.001}}</ref><ref>{{Cite journal |last=Tlak Gajger |first=Ivana |last2=Šimenc |first2=Laura |last3=Toplak |first3=Ivan |date=2021-06-25 |title=The First Detection and Genetic Characterization of Four Different Honeybee Viruses in Wild Bumblebees from Croatia |url=http://dx.doi.org/10.3390/pathogens10070808 |journal=Pathogens |volume=10 |issue=7 |pages=808 |doi=10.3390/pathogens10070808 |issn=2076-0817}}</ref>


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|Hepatitis E
|Hepatitis E
|Wild Boar<ref>{{Cite journal |last1=Anheyer-Behmenburg |first1=Helena E. |last2=Szabo |first2=Kathrin |last3=Schotte |first3=Ulrich |last4=Binder |first4=Alfred |last5=Klein |first5=Günter |last6=Johne |first6=Reimar |title=Hepatitis E Virus in Wild Boars and Spillover Infection in Red and Roe Deer, Germany, 2013–2015 |journal=Emerging Infectious Diseases |year=2017 |volume=23 |issue=1 |pages=130–133 |url=https://wwwnc.cdc.gov/eid/article/23/1/16-1169_article |language=en-us |doi=10.3201/eid2301.161169|pmid=27983488 |pmc=5176221 }}</ref>
|Wild Boar<ref name=":3">{{Cite journal |last1=Anheyer-Behmenburg |first1=Helena E. |last2=Szabo |first2=Kathrin |last3=Schotte |first3=Ulrich |last4=Binder |first4=Alfred |last5=Klein |first5=Günter |last6=Johne |first6=Reimar |title=Hepatitis E Virus in Wild Boars and Spillover Infection in Red and Roe Deer, Germany, 2013–2015 |journal=Emerging Infectious Diseases |year=2017 |volume=23 |issue=1 |pages=130–133 |url=https://wwwnc.cdc.gov/eid/article/23/1/16-1169_article |language=en-us |doi=10.3201/eid2301.161169|pmid=27983488 |pmc=5176221 }}</ref>
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|Ebola
|Ebola
|Fruit Bats<ref>{{cite journal | doi=10.1371/journal.pone.0271886 | doi-access=free | title=Estimation of Ebola's spillover infection exposure in Sierra Leone based on sociodemographic and economic factors | year=2022 | last1=Mursel | first1=Sena | last2=Alter | first2=Nathaniel | last3=Slavit | first3=Lindsay | last4=Smith | first4=Anna | last5=Bocchini | first5=Paolo | last6=Buceta | first6=Javier | journal=PLOS ONE | volume=17 | issue=9 | pages=e0271886 | pmid=36048780 | pmc=9436100 | arxiv=2109.15313 | bibcode=2022PLoSO..1771886M }}</ref>
|Fruit Bats<ref name=":4">{{cite journal | doi=10.1371/journal.pone.0271886 | doi-access=free | title=Estimation of Ebola's spillover infection exposure in Sierra Leone based on sociodemographic and economic factors | year=2022 | last1=Mursel | first1=Sena | last2=Alter | first2=Nathaniel | last3=Slavit | first3=Lindsay | last4=Smith | first4=Anna | last5=Bocchini | first5=Paolo | last6=Buceta | first6=Javier | journal=PLOS ONE | volume=17 | issue=9 | pages=e0271886 | pmid=36048780 | pmc=9436100 | arxiv=2109.15313 | bibcode=2022PLoSO..1771886M }}</ref>
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|HIV/AIDS
|HIV/AIDS
|Chimpanzee<ref>{{Cite web |date=2022-06-30 |title=About HIV/AIDS {{!}} HIV Basics {{!}} HIV/AIDS {{!}} CDC |url=https://www.cdc.gov/hiv/basics/whatishiv.html |access-date=2022-12-08 |website=www.cdc.gov |language=en-us}}</ref>
|Chimpanzee<ref name=":5">{{Cite web |date=2022-06-30 |title=About HIV/AIDS {{!}} HIV Basics {{!}} HIV/AIDS {{!}} CDC |url=https://www.cdc.gov/hiv/basics/whatishiv.html |access-date=2022-12-08 |website=www.cdc.gov |language=en-us}}</ref>
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|COVID-19
|COVID-19
|Bats{{refn|name=COVID_Reservoir|SARS-CoV-2 is widely believed to have an original reservoir in bats,<ref>{{Cite journal |last1=Valencak |first1=Teresa G. |last2=Csiszar |first2=Anna |last3=Szalai |first3=Gabor |last4=Podlutsky |first4=Andrej |last5=Tarantini |first5=Stefano |last6=Fazekas-Pongor |first6=Vince |last7=Papp |first7=Magor |last8=Ungvari |first8=Zoltan |date=October 2021 |title=Animal reservoirs of SARS-CoV-2: calculable COVID-19 risk for older adults from animal to human transmission |journal=GeroScience |volume=43 |issue=5 |pages=2305–2320 |doi=10.1007/s11357-021-00444-9 |issn=2509-2723 |pmc=8404404 |pmid=34460063}}</ref><ref name="Lawler2021">{{cite journal |last1=Lawler |first1=Odette K |last2=Allan |first2=Hannah L |last3=Baxter |first3=Peter W J |last4=Castagnino |first4=Romi |last5=Tor |first5=Marina Corella |last6=Dann |first6=Leah E |last7=Hungerford |first7=Joshua |last8=Karmacharya |first8=Dibesh |last9=Lloyd |first9=Thomas J |last10=López-Jara |first10=María José |last11=Massie |first11=Gloeta N |last12=Novera |first12=Junior |last13=Rogers |first13=Andrew M |last14=Kark |first14=Salit |title=The COVID-19 pandemic is intricately linked to biodiversity loss and ecosystem health |journal=The Lancet. Planetary Health |date=November 2021 |volume=5 |issue=11 |pages=e840–e850 |doi=10.1016/S2542-5196(21)00258-8 |pmid=34774124 |pmc=8580505 |quote=The current weight of evidence suggests that SARS-CoV-2, or its progenitor, probably emerged in humans from a zoonotic source in Wuhan, China, where it was first identified in 2019. Although evidence on the origins of SARS-CoV-2 are inconclusive, bats have been suggested to be the most probable evolutionary source for the virus."}}</ref><ref name="Castelo-BrancoNobreSouza2023">{{cite journal | last1 = Castelo-Branco | first1 = D.S.C.M. | last2 = Nobre | first2 = J.A. | last3 = Souza | first3 = P.R.H. | last4 = Diógenes | first4 = E.M. | last5 = Guedes | first5 = G.M.M. | last6 = Mesquita | first6 = F.P. | last7 = Souza | first7 = P.F.N. | last8 = Rocha | first8 = M.F.G. | last9 = Sidrim | first9 = J.J.C. | last10 = Cordeiro | first10 = R.A. | last11 = Montenegro | first11 = R.C. | title = Role of Brazilian bats in the epidemiological cycle of potentially zoonotic pathogens | journal = Microbial Pathogenesis | date = February 2023 | volume = 177 | page = 106032 | issn = 0882-4010 | doi = 10.1016/j.micpath.2023.106032 | pmid = 36804526| s2cid = 257015965 | url = |quote="The pandemic of Coronavirus disease (COVID-19) has highlighted bats as reservoirs of coronaviruses that cause severe respiratory diseases in humans and, frequently, in other animals. However, despite the spillover events of SARS-CoV and MERS-CoV, the implication of bats as natural reservoirs of the ancient virus of SARS-CoV-2 is, to date, unconfirmed, as only closely related SARS-like viruses have been detected by genomic sequencing and little is known about the mechanisms of host switch from bats to humans."}}</ref> though there may have been an intermediate host (such as [[palm civet]]s,<ref name="YuanJiangLi2020">{{cite journal | last1 = Yuan | first1 = Shu | last2 = Jiang | first2 = Si-Cong | last3 = Li | first3 = Zi-Lin | title = Analysis of Possible Intermediate Hosts of the New Coronavirus SARS-CoV-2 | journal = Frontiers in Veterinary Science | date = 9 June 2020 | volume = 7 | page = 379 | eissn = 2297-1769 | doi = 10.3389/fvets.2020.00379 | pmid = 32582786 | pmc = 7297130 | url = | doi-access = free }}</ref><ref name="ZhouShi2021"/> [[minks]],<ref name="Oude MunninkSikkemaNieuwenhuijse2021">{{cite journal | last1 = Oude Munnink | first1 = Bas B. | last2 = Sikkema | first2 = Reina S. | last3 = Nieuwenhuijse | first3 = David F. | last4 = Molenaar | first4 = Robert Jan | last5 = Munger | first5 = Emmanuelle | last6 = Molenkamp | first6 = Richard | last7 = van der Spek | first7 = Arco | last8 = Tolsma | first8 = Paulien | last9 = Rietveld | first9 = Ariene | last10 = Brouwer | first10 = Miranda | last11 = Bouwmeester-Vincken | first11 = Noortje | last12 = Harders | first12 = Frank | last13 = Hakze-van der Honing | first13 = Renate | last14 = Wegdam-Blans | first14 = Marjolein C. A. | last15 = Bouwstra | first15 = Ruth J. | last16 = GeurtsvanKessel | first16 = Corine | last17 = van der Eijk | first17 = Annemiek A. | last18 = Velkers | first18 = Francisca C. | last19 = Smit | first19 = Lidwien A. M. | last20 = Stegeman | first20 = Arjan | last21 = van der Poel | first21 = Wim H. M. | last22 = Koopmans | first22 = Marion P. G. | title = Transmission of SARS-CoV-2 on mink farms between humans and mink and back to humans | journal = Science | date = 8 January 2021 | volume = 371 | issue = 6525 | pages = 172–177 | issn = 0036-8075 | eissn = 1095-9203 | doi = 10.1126/science.abe5901 | pmid = 33172935 | pmc = 7857398 | bibcode = 2021Sci...371..172O | url = }}</ref><ref name="ZhouShi2021">{{cite journal | last1 = Zhou | first1 = Peng | last2 = Shi | first2 = Zheng-Li | title = SARS-CoV-2 spillover events | journal = Science | date = 8 January 2021 | volume = 371 | issue = 6525 | pages = 120–122 | issn = 0036-8075 | eissn = 1095-9203 | doi = 10.1126/science.abf6097 | pmid = 33414206 | bibcode = 2021Sci...371..120Z | s2cid = 231138544 | url = | doi-access = free }}</ref> or [[pangolins]]<ref name="SinghYi2021">{{cite journal | last1 = Singh | first1 = Devika | last2 = Yi | first2 = Soojin V. | title = On the origin and evolution of SARS-CoV-2 | journal = Experimental & Molecular Medicine | date = April 2021 | volume = 53 | issue = 4 | pages = 537–547 | issn = 1226-3613 | eissn = 2092-6413 | doi = 10.1038/s12276-021-00604-z | pmid = 33864026 | pmc = 8050477 | url = }}</ref><ref name="WrobelBentonXu2021">{{cite journal | last1 = Wrobel | first1 = Antoni G. | last2 = Benton | first2 = Donald J. | last3 = Xu | first3 = Pengqi | last4 = Calder | first4 = Lesley J. | last5 = Borg | first5 = Annabel | last6 = Roustan | first6 = Chloë | last7 = Martin | first7 = Stephen R. | last8 = Rosenthal | first8 = Peter B. | last9 = Skehel | first9 = John J. | last10 = Gamblin | first10 = Steven J. | title = Structure and binding properties of Pangolin-CoV spike glycoprotein inform the evolution of SARS-CoV-2 | journal = Nature Communications | date = 5 February 2021 | volume = 12 | issue = 1 | page = 837 | eissn = 2041-1723 | doi = 10.1038/s41467-021-21006-9 | pmid = 33547281 | pmc = 7864994 | bibcode = 2021NatCo..12..837W | url = }}</ref>) before spillover into humans.<ref name="PerlmanPeiris2023">{{cite journal | last1 = Perlman | first1 = Stanley | last2 = Peiris | first2 = Malik | title = Coronavirus research: knowledge gaps and research priorities | journal = Nature Reviews Microbiology | date = 15 February 2023 | volume = 21 | issue = 3 | pages = 125–126 | issn = 1740-1526 | eissn = 1740-1534 | doi = 10.1038/s41579-022-00837-3 | pmid = 36792727 | s2cid = 256875846 | url = |quote="It is almost certain that the virus originated in bats and crossed species to humans either directly or indirectly via intermediary hosts."| doi-access = free }}</ref><ref name="Keusch2022">{{cite journal |last1=Keusch |first1=Gerald T. |last2=Amuasi |first2=John H. |last3=Anderson |first3=Danielle E. |last4=Daszak |first4=Peter |last5=Eckerle |first5=Isabella |last6=Field |first6=Hume |last7=Koopmans |first7=Marion |last8=Lam |first8=Sai Kit |last9=Das Neves |first9=Carlos G. |last10=Peiris |first10=Malik |last11=Perlman |first11=Stanley |last12=Wacharapluesadee |first12=Supaporn |last13=Yadana |first13=Su |last14=Saif |first14=Linda |title=Pandemic origins and a One Health approach to preparedness and prevention: Solutions based on SARS-CoV-2 and other RNA viruses |journal=Proceedings of the National Academy of Sciences |date=18 October 2022 |volume=119 |issue=42 |pages=e2202871119 |doi=10.1073/pnas.2202871119 |pmid=36215506 |pmc=9586299 |bibcode=2022PNAS..11902871K |s2cid=252818019 |language=en |issn=0027-8424|quote="The increasing scientific evidence concerning the origins of Severe Acute Respiratory Syndrome Coronavirus-2 (SARS-CoV-2) is most consistent with a zoonotic origin and a spillover pathway from wildlife to people via wildlife farming and the wildlife trade."}}</ref>}}
|Bats{{refn|name=COVID_Reservoir|SARS-CoV-2 is widely believed to have an original reservoir in bats,<ref>{{Cite journal |last1=Valencak |first1=Teresa G. |last2=Csiszar |first2=Anna |last3=Szalai |first3=Gabor |last4=Podlutsky |first4=Andrej |last5=Tarantini |first5=Stefano |last6=Fazekas-Pongor |first6=Vince |last7=Papp |first7=Magor |last8=Ungvari |first8=Zoltan |date=October 2021 |title=Animal reservoirs of SARS-CoV-2: calculable COVID-19 risk for older adults from animal to human transmission |journal=GeroScience |volume=43 |issue=5 |pages=2305–2320 |doi=10.1007/s11357-021-00444-9 |issn=2509-2723 |pmc=8404404 |pmid=34460063}}</ref><ref name="Lawler2021">{{cite journal |last1=Lawler |first1=Odette K |last2=Allan |first2=Hannah L |last3=Baxter |first3=Peter W J |last4=Castagnino |first4=Romi |last5=Tor |first5=Marina Corella |last6=Dann |first6=Leah E |last7=Hungerford |first7=Joshua |last8=Karmacharya |first8=Dibesh |last9=Lloyd |first9=Thomas J |last10=López-Jara |first10=María José |last11=Massie |first11=Gloeta N |last12=Novera |first12=Junior |last13=Rogers |first13=Andrew M |last14=Kark |first14=Salit |title=The COVID-19 pandemic is intricately linked to biodiversity loss and ecosystem health |journal=The Lancet. Planetary Health |date=November 2021 |volume=5 |issue=11 |pages=e840–e850 |doi=10.1016/S2542-5196(21)00258-8 |pmid=34774124 |pmc=8580505 |quote=The current weight of evidence suggests that SARS-CoV-2, or its progenitor, probably emerged in humans from a zoonotic source in Wuhan, China, where it was first identified in 2019. Although evidence on the origins of SARS-CoV-2 are inconclusive, bats have been suggested to be the most probable evolutionary source for the virus."}}</ref><ref name="Castelo-BrancoNobreSouza2023">{{cite journal | last1 = Castelo-Branco | first1 = D.S.C.M. | last2 = Nobre | first2 = J.A. | last3 = Souza | first3 = P.R.H. | last4 = Diógenes | first4 = E.M. | last5 = Guedes | first5 = G.M.M. | last6 = Mesquita | first6 = F.P. | last7 = Souza | first7 = P.F.N. | last8 = Rocha | first8 = M.F.G. | last9 = Sidrim | first9 = J.J.C. | last10 = Cordeiro | first10 = R.A. | last11 = Montenegro | first11 = R.C. | title = Role of Brazilian bats in the epidemiological cycle of potentially zoonotic pathogens | journal = Microbial Pathogenesis | date = February 2023 | volume = 177 | page = 106032 | issn = 0882-4010 | doi = 10.1016/j.micpath.2023.106032 | pmid = 36804526| s2cid = 257015965 | url = |quote="The pandemic of Coronavirus disease (COVID-19) has highlighted bats as reservoirs of coronaviruses that cause severe respiratory diseases in humans and, frequently, in other animals. However, despite the spillover events of SARS-CoV and MERS-CoV, the implication of bats as natural reservoirs of the ancient virus of SARS-CoV-2 is, to date, unconfirmed, as only closely related SARS-like viruses have been detected by genomic sequencing and little is known about the mechanisms of host switch from bats to humans."}}</ref> though there may have been an intermediate host (such as [[palm civet]]s,<ref name="YuanJiangLi2020">{{cite journal | last1 = Yuan | first1 = Shu | last2 = Jiang | first2 = Si-Cong | last3 = Li | first3 = Zi-Lin | title = Analysis of Possible Intermediate Hosts of the New Coronavirus SARS-CoV-2 | journal = Frontiers in Veterinary Science | date = 9 June 2020 | volume = 7 | page = 379 | eissn = 2297-1769 | doi = 10.3389/fvets.2020.00379 | pmid = 32582786 | pmc = 7297130 | url = | doi-access = free }}</ref><ref name="ZhouShi2021"/> [[minks]],<ref name="Oude MunninkSikkemaNieuwenhuijse2021">{{cite journal | last1 = Oude Munnink | first1 = Bas B. | last2 = Sikkema | first2 = Reina S. | last3 = Nieuwenhuijse | first3 = David F. | last4 = Molenaar | first4 = Robert Jan | last5 = Munger | first5 = Emmanuelle | last6 = Molenkamp | first6 = Richard | last7 = van der Spek | first7 = Arco | last8 = Tolsma | first8 = Paulien | last9 = Rietveld | first9 = Ariene | last10 = Brouwer | first10 = Miranda | last11 = Bouwmeester-Vincken | first11 = Noortje | last12 = Harders | first12 = Frank | last13 = Hakze-van der Honing | first13 = Renate | last14 = Wegdam-Blans | first14 = Marjolein C. A. | last15 = Bouwstra | first15 = Ruth J. | last16 = GeurtsvanKessel | first16 = Corine | last17 = van der Eijk | first17 = Annemiek A. | last18 = Velkers | first18 = Francisca C. | last19 = Smit | first19 = Lidwien A. M. | last20 = Stegeman | first20 = Arjan | last21 = van der Poel | first21 = Wim H. M. | last22 = Koopmans | first22 = Marion P. G. | title = Transmission of SARS-CoV-2 on mink farms between humans and mink and back to humans | journal = Science | date = 8 January 2021 | volume = 371 | issue = 6525 | pages = 172–177 | issn = 0036-8075 | eissn = 1095-9203 | doi = 10.1126/science.abe5901 | pmid = 33172935 | pmc = 7857398 | bibcode = 2021Sci...371..172O | url = }}</ref><ref name="ZhouShi2021">{{cite journal | last1 = Zhou | first1 = Peng | last2 = Shi | first2 = Zheng-Li | title = SARS-CoV-2 spillover events | journal = Science | date = 8 January 2021 | volume = 371 | issue = 6525 | pages = 120–122 | issn = 0036-8075 | eissn = 1095-9203 | doi = 10.1126/science.abf6097 | pmid = 33414206 | bibcode = 2021Sci...371..120Z | s2cid = 231138544 | url = | doi-access = free }}</ref> or [[pangolins]]<ref name="SinghYi2021">{{cite journal | last1 = Singh | first1 = Devika | last2 = Yi | first2 = Soojin V. | title = On the origin and evolution of SARS-CoV-2 | journal = Experimental & Molecular Medicine | date = April 2021 | volume = 53 | issue = 4 | pages = 537–547 | issn = 1226-3613 | eissn = 2092-6413 | doi = 10.1038/s12276-021-00604-z | pmid = 33864026 | pmc = 8050477 | url = }}</ref><ref name="WrobelBentonXu2021">{{cite journal | last1 = Wrobel | first1 = Antoni G. | last2 = Benton | first2 = Donald J. | last3 = Xu | first3 = Pengqi | last4 = Calder | first4 = Lesley J. | last5 = Borg | first5 = Annabel | last6 = Roustan | first6 = Chloë | last7 = Martin | first7 = Stephen R. | last8 = Rosenthal | first8 = Peter B. | last9 = Skehel | first9 = John J. | last10 = Gamblin | first10 = Steven J. | title = Structure and binding properties of Pangolin-CoV spike glycoprotein inform the evolution of SARS-CoV-2 | journal = Nature Communications | date = 5 February 2021 | volume = 12 | issue = 1 | page = 837 | eissn = 2041-1723 | doi = 10.1038/s41467-021-21006-9 | pmid = 33547281 | pmc = 7864994 | bibcode = 2021NatCo..12..837W | url = }}</ref>) before spillover into humans.<ref name="PerlmanPeiris2023">{{cite journal | last1 = Perlman | first1 = Stanley | last2 = Peiris | first2 = Malik | title = Coronavirus research: knowledge gaps and research priorities | journal = Nature Reviews Microbiology | date = 15 February 2023 | volume = 21 | issue = 3 | pages = 125–126 | issn = 1740-1526 | eissn = 1740-1534 | doi = 10.1038/s41579-022-00837-3 | pmid = 36792727 | s2cid = 256875846 | url = |quote="It is almost certain that the virus originated in bats and crossed species to humans either directly or indirectly via intermediary hosts."| doi-access = free }}</ref><ref name="Keusch2022">{{cite journal |last1=Keusch |first1=Gerald T. |last2=Amuasi |first2=John H. |last3=Anderson |first3=Danielle E. |last4=Daszak |first4=Peter |last5=Eckerle |first5=Isabella |last6=Field |first6=Hume |last7=Koopmans |first7=Marion |last8=Lam |first8=Sai Kit |last9=Das Neves |first9=Carlos G. |last10=Peiris |first10=Malik |last11=Perlman |first11=Stanley |last12=Wacharapluesadee |first12=Supaporn |last13=Yadana |first13=Su |last14=Saif |first14=Linda |title=Pandemic origins and a One Health approach to preparedness and prevention: Solutions based on SARS-CoV-2 and other RNA viruses |journal=Proceedings of the National Academy of Sciences |date=18 October 2022 |volume=119 |issue=42 |pages=e2202871119 |doi=10.1073/pnas.2202871119 |pmid=36215506 |pmc=9586299 |bibcode=2022PNAS..11902871K |s2cid=252818019 |language=en |issn=0027-8424|quote="The increasing scientific evidence concerning the origins of Severe Acute Respiratory Syndrome Coronavirus-2 (SARS-CoV-2) is most consistent with a zoonotic origin and a spillover pathway from wildlife to people via wildlife farming and the wildlife trade."}}</ref>}}
|}
|}

== Causes of Spillover ==
Zoonotic spillover is a relatively uncommon but incredibly dangerous natural phenomenon--as is evidenced by the Ebola epidemic and Coronavirus pandemic. For zoonotic spillover to occur, several important factors have to occur in tandem.<ref name=":2" /> Such factors include altered ecological niches, epidemiological susceptibility, and the natural behavior of pathogens and novel host or spillover host species.<ref>{{Cite journal |last=Walsh |first=Michael G. |last2=Wiethoelter |first2=Anke |last3=Haseeb |first3=M. A. |date=2017-08-15 |title=The impact of human population pressure on flying fox niches and the potential consequences for Hendra virus spillover |url=https://www.nature.com/articles/s41598-017-08065-z |journal=Scientific Reports |language=en |volume=7 |issue=1 |doi=10.1038/s41598-017-08065-z |issn=2045-2322}}</ref> By suggesting that the natural behavior of pathogens and host species impacts zoonotic spillover, simple Darwinian theories are being referenced. As with all species, a pathogen's main goal is to survive. When a stressor puts pressure on the survival of the pathogenic species, it will have to adapt to said stressor in order to survive.<ref name=":6">{{Cite journal |last=Becker |first=Daniel J. |last2=Eby |first2=Peggy |last3=Madden |first3=Wyatt |last4=Peel |first4=Alison J. |last5=Plowright |first5=Raina K. |date=2023-01 |editor-last=Ostfeld |editor-first=Richard |title=Ecological conditions predict the intensity of Hendra virus excretion over space and time from bat reservoir hosts |url=https://onlinelibrary.wiley.com/doi/10.1111/ele.14007 |journal=Ecology Letters |language=en |volume=26 |issue=1 |pages=23–36 |doi=10.1111/ele.14007 |issn=1461-023X}}</ref> For example, the ecological niche of the novel host may be subject to a lack of food which leads to a decrease in the novel host population. In order for a virus to replicate, it must invade a eukaryotic organism.<ref>{{Citation |last=Louten |first=Jennifer |title=Virus Replication |date=2016 |url=https://linkinghub.elsevier.com/retrieve/pii/B9780128009475000041 |work=Essential Human Virology |pages=49–70 |access-date=2023-10-26 |publisher=Elsevier |language=en |doi=10.1016/b978-0-12-800947-5.00004-1. pmc 7149683. |isbn=978-0-12-800947-5}}</ref> When the novel eukaryotic organism is not available for the virus to infect, it must jump to another host.<ref name=":6" /> In order for the virus to make the jump to the spillover host, the spillover host must be epidemiologically susceptible to this virus. Although it is not well understood what makes one spillover host "better" than another host, it is known that the susceptibility has to do with the shedding rate of the virus, how well the virus survives and moves while not within a host, the genotypic similarities between the novel and spillover hosts, and the behavior of the spillover host that leads to contact with a high dose of the virus.<ref name=":2" />


== See also ==
== See also ==

Revision as of 20:26, 26 October 2023

Spillover infection, also known as pathogen spillover and spillover event, occurs when a reservoir population with a high pathogen prevalence comes into contact with a novel host population. The pathogen is transmitted from the reservoir population and may or may not be transmitted within the host population.[1] Due to climate change and land use expansion, the risk of viral spillover is predicted to significantly increase.[2][3]

Spillover zoonoses

The fruit bat is believed to be the zoonotic agent responsible for the spillover of the Ebola virus.

Spillover is a common event; in fact, more than two-thirds of human viruses are zoonotic.[4][5] Most spillover events result in self-limited cases with no further human-to-human transmission, as occurs, for example, with rabies, anthrax, histoplasmosis or hydatidosis. Other zoonotic pathogens are able to be transmitted by humans to produce secondary cases and even to establish limited chains of transmission. Some examples are the Ebola and Marburg filoviruses, the MERS and SARS coronaviruses and some avian flu viruses. Finally, some spillover events can result in the final adaptation of the microbe to humans, who can become a new stable reservoir, as occurred with the HIV virus resulting in the AIDS epidemic and with SARS-CoV-2 resulting in the COVID-19 pandemic.[5]

If the history of mutual adaptation is long enough, permanent host-microbe associations can be established resulting in co-evolution, and even permanent integration of the microbe genome with the human genome, as is the case of endogenous viruses.[6] The closer the two target host species are in phylogenetic terms, the easier it is for microbes to overcome the biological barrier to produce successful spillovers.[1] For this reason, other mammals are the main source of zoonotic agents for humans. For example, in the case of the Ebola virus, fruit bats are the hypothesized zoonotic agent.[7]

During the late 20th century, zoonotic spillover increased as the environmental impact of agriculture promoted increased land use and deforestation, changing wildlife habitat. As species shift their geographic range in response to climate change, the risk of zoonotic spillover is predicted to substantially increase, particularly in tropical regions that are experiencing rapid warming.[8] As forested areas of land are cleared for human use, there is increased proximity and interaction between wild animals and humans thereby increasing the potential for exposure.[9]

Intraspecies spillover

The bumblebee is a potential reservoir for several pollinator parasites.

Commercially bred bumblebees used to pollinate greenhouses can be reservoirs for several pollinator parasites including the protozoans Crithidia bombi, and Apicystis bombi,[10] the microsporidians Nosema bombi and Nosema ceranae,[10][11] plus viruses such as Deformed wing virus and the tracheal mites Locustacarus buchneri.[11] Commercial bees that escape the greenhouse environment may then infect wild bee populations. Infection may be via direct interactions between managed and wild bees or via shared flower use and contamination.[12][13] One study found that half of all wild bees found near greenhouses were infected with C. bombi. Rates and incidence of infection decline dramatically the further away from the greenhouses where the wild bees are located.[14][15] Instances of spillover between bumblebees are well documented across the world, particularly in Japan, North America, and the United Kingdom.[16][17]

Examples of Spillover Zoonosis
Disease Reservoir
Hepatitis E Wild Boar[10]
Ebola Fruit Bats[11]
HIV/AIDS Chimpanzee[12]
COVID-19 Bats[28]

Causes of Spillover

Zoonotic spillover is a relatively uncommon but incredibly dangerous natural phenomenon--as is evidenced by the Ebola epidemic and Coronavirus pandemic. For zoonotic spillover to occur, several important factors have to occur in tandem.[1] Such factors include altered ecological niches, epidemiological susceptibility, and the natural behavior of pathogens and novel host or spillover host species.[29] By suggesting that the natural behavior of pathogens and host species impacts zoonotic spillover, simple Darwinian theories are being referenced. As with all species, a pathogen's main goal is to survive. When a stressor puts pressure on the survival of the pathogenic species, it will have to adapt to said stressor in order to survive.[30] For example, the ecological niche of the novel host may be subject to a lack of food which leads to a decrease in the novel host population. In order for a virus to replicate, it must invade a eukaryotic organism.[31] When the novel eukaryotic organism is not available for the virus to infect, it must jump to another host.[30] In order for the virus to make the jump to the spillover host, the spillover host must be epidemiologically susceptible to this virus. Although it is not well understood what makes one spillover host "better" than another host, it is known that the susceptibility has to do with the shedding rate of the virus, how well the virus survives and moves while not within a host, the genotypic similarities between the novel and spillover hosts, and the behavior of the spillover host that leads to contact with a high dose of the virus.[1]

See also

References

  1. ^ a b c d Woolhouse, Mark; Scott, Fiona; Hudson, Zoe; Howey, Richard; Chase-Topping, Margo (2012). "Human viruses: Discovery and emergence". Philosophical Transactions of the Royal Society B: Biological Sciences. 367 (1604): 2864–2871. doi:10.1098/rstb.2011.0354. PMC 3427559. PMID 22966141.
  2. ^ Wolfe, Nathan D.; Dunavan, Claire Panosian; Diamond, Jared (May 2007). "Origins of major human infectious diseases". Nature. 447 (7142): 279–283. Bibcode:2007Natur.447..279W. doi:10.1038/nature05775. ISSN 1476-4687. PMC 7095142. PMID 17507975.
  3. ^ Ebola. (2014). National Center for Emerging and Zoonotic Infectious Diseases, Division of High-Consequence Pathogens and Pathology, Department of Health & Human Services, CDC.
  4. ^ Graystock, P; Yates, K; Evison, SEF; Darvill, B; Goulson, D; Hughes, WOH (2013). "The Trojan hives: pollinator pathogens, imported and distributed in bumblebee colonies". Journal of Applied Ecology. 50 (5): 1207–15. doi:10.1111/1365-2664.12134. S2CID 3937352.
  5. ^ a b Sachman-Ruiz, Bernardo; Narváez-Padilla, Verónica; Reynaud, Enrique (2015-03-10). "Commercial Bombus impatiens as reservoirs of emerging infectious diseases in central México". Biological Invasions. 17 (7): 2043–53. doi:10.1007/s10530-015-0859-6. ISSN 1387-3547.
  6. ^ Durrer, Stephan; Schmid-Hempel, Paul (1994-12-22). "Shared Use of Flowers Leads to Horizontal Pathogen Transmission". Proceedings of the Royal Society of London B: Biological Sciences. 258 (1353): 299–302. Bibcode:1994RSPSB.258..299D. doi:10.1098/rspb.1994.0176. ISSN 0962-8452. S2CID 84926310.
  7. ^ Graystock, Peter; Goulson, Dave; Hughes, William O. H. (2015-08-22). "Parasites in bloom: flowers aid dispersal and transmission of pollinator parasites within and between bee species". Proceedings of the Royal Society B: Biological Sciences. 282 (1813): 20151371. doi:10.1098/rspb.2015.1371. ISSN 0962-8452. PMC 4632632. PMID 26246556.
  8. ^ Otterstatter, MC; Thomson, JD (2008). "Does Pathogen Spillover from Commercially Reared Bumble Bees Threaten Wild Pollinators?". PLOS ONE. 3 (7): e2771. Bibcode:2008PLoSO...3.2771O. doi:10.1371/journal.pone.0002771. PMC 2464710. PMID 18648661.
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