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Viruses are an important natural means of [[Horizontal gene transfer|transferring genes]] between different species, which increases [[genetic diversity]] and drives evolution.<ref name="Canchaya">{{vcite journal|author=Canchaya C, Fournous G, Chibani-Chennoufi S, Dillmann ML, Brüssow H|title=Phage as agents of lateral gene transfer|journal=Current Opinion in Microbiology |volume=6 |issue=4 |pages=417–24 |year=2003 |pmid=12941415|doi=10.1016/S1369-5274(03)00086-9}}</ref> It is thought that viruses played a central role in the early evolution, before the diversification of bacteria, archaea and eukaryotes, at the time of the [[Last universal ancestor|last universal common ancestor]] of life on Earth.<ref name="pmid11536914">{{vcite journal |author=Forterre P, Philippe H |title=The last universal common ancestor (LUCA), simple or complex? |journal=The Biological Bulletin |volume=196 |issue=3 |pages=373–5; discussion 375–7 |year=1999 |pmid=11536914 |doi= 10.2307/1542973}}</ref> Viruses are still one of the largest reservoirs of unexplored genetic diversity on Earth.<ref name="pmid17853907" />
Viruses are an important natural means of [[Horizontal gene transfer|transferring genes]] between different species, which increases [[genetic diversity]] and drives evolution.<ref name="Canchaya">{{vcite journal|author=Canchaya C, Fournous G, Chibani-Chennoufi S, Dillmann ML, Brüssow H|title=Phage as agents of lateral gene transfer|journal=Current Opinion in Microbiology |volume=6 |issue=4 |pages=417–24 |year=2003 |pmid=12941415|doi=10.1016/S1369-5274(03)00086-9}}</ref> It is thought that viruses played a central role in the early evolution, before the diversification of bacteria, archaea and eukaryotes, at the time of the [[Last universal ancestor|last universal common ancestor]] of life on Earth.<ref name="pmid11536914">{{vcite journal |author=Forterre P, Philippe H |title=The last universal common ancestor (LUCA), simple or complex? |journal=The Biological Bulletin |volume=196 |issue=3 |pages=373–5; discussion 375–7 |year=1999 |pmid=11536914 |doi= 10.2307/1542973}}</ref> Viruses are still one of the largest reservoirs of unexplored genetic diversity on Earth.<ref name="pmid17853907" />

[[Virus]]es are part of the [[hydrothermal vent microbial community]] and their influence on the microbial ecology in these ecosystems is a burgeoning field of research.<ref>{{Cite journal|last=Anderson|first=Rika E.|last2=Brazelton|first2=William J.|last3=Baross|first3=John A.|date=2011|title=Is the genetic landscape of the deep subsurface biosphere affected by viruses?|journal=Frontiers in Microbiology|volume=2|pages=219|doi=10.3389/fmicb.2011.00219|issn=1664-302X|pmc=3211056|pmid=22084639}}</ref> Viruses are the most abundant life in the ocean, harboring the greatest reservoir of genetic diversity.<ref>{{Cite journal|last=Suttle|first=Curtis A.|date=September 2005|title=Viruses in the sea|journal=Nature|volume=437|issue=7057|pages=356–361|doi=10.1038/nature04160|pmid=16163346|issn=0028-0836|bibcode=2005Natur.437..356S}}</ref> As their infections are often fatal, they constitute a significant source of mortality and thus have widespread influence on biological oceanographic processes, [[evolution]] and [[Geochemical cycle#Differentiating biogeochemical cycles|biogeochemical cycling]] within the ocean.<ref>{{Cite journal|last=Suttle|first=Curtis A.|date=October 2007|title=Marine viruses — major players in the global ecosystem|journal=Nature Reviews Microbiology|language=En|volume=5|issue=10|pages=801–812|doi=10.1038/nrmicro1750|pmid=17853907|issn=1740-1526}}</ref> Evidence has been found however to indicate that viruses found in vent habitats have adopted a more [[Mutualism (biology)|mutualistic]] than [[Parasitism|parasitic]] evolutionary strategy in order to survive the extreme and volatile environment they exist in.<ref name=":17">{{Cite journal|last=Anderson|first=Rika E.|last2=Sogin|first2=Mitchell L.|last3=Baross|first3=John A.|date=2014-10-03|title=Evolutionary Strategies of Viruses, Bacteria and Archaea in Hydrothermal Vent Ecosystems Revealed through Metagenomics|journal=PLOS ONE|volume=9|issue=10|pages=e109696|doi=10.1371/journal.pone.0109696|issn=1932-6203|pmc=4184897|pmid=25279954|bibcode=2014PLoSO...9j9696A}}</ref> Deep-sea hydrothermal vents were found to have high numbers of viruses indicating high viral production.<ref name=":18">{{Cite journal|last=Ortmann|first=Alice C.|last2=Suttle|first2=Curtis A.|date=August 2005|title=High abundances of viruses in a deep-sea hydrothermal vent system indicates viral mediated microbial mortality|journal=Deep Sea Research Part I: Oceanographic Research Papers|volume=52|issue=8|pages=1515–1527|doi=10.1016/j.dsr.2005.04.002|issn=0967-0637|bibcode=2005DSRI...52.1515O}}</ref> Like in other marine environments, [[Viruses and deep-sea hydrothermal vents|deep-sea hydrothermal viruses]] affect abundance and diversity of [[prokaryote]]s and therefore impact microbial biogeochemical cycling by [[Lysis|lysing]] their hosts to replicate.<ref>{{Cite journal|authorlink=Mya Breitbart|last=Breitbart|first=Mya|date=2012-01-15|title=Marine Viruses: Truth or Dare|journal=Annual Review of Marine Science|language=en|volume=4|issue=1|pages=425–448|doi=10.1146/annurev-marine-120709-142805|pmid=22457982|issn=1941-1405|bibcode=2012ARMS....4..425B}}</ref> However, in contrast to their role as a source of mortality and population control, viruses have also been postulated to enhance survival of prokaryotes in extreme environments, acting as reservoirs of genetic information. The interactions of the virosphere with microorganisms under environmental stresses is therefore thought to aide microorganism survival through dispersal of host genes through [[Horizontal gene transfer in evolution|horizontal gene transfer]].<ref>{{Cite journal|last=Goldenfeld|first=Nigel|last2=Woese|first2=Carl|date=January 2007|title=Biology's next revolution|journal=Nature|volume=445|issue=7126|pages=369|doi=10.1038/445369a|pmid=17251963|issn=0028-0836|arxiv=q-b


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Revision as of 09:57, 10 May 2020

Virus-host interactions in the marine ecosystem,
including viral infection of bacteria, phytoplankton and fish[1]

A virus is a small infectious agent that replicates only inside the living cells of other organisms. Viruses can infect all types of life forms, from animals and plants to microorganisms, including bacteria and archaea.[2]

When not inside an infected cell or in the process of infecting a cell, viruses exist in the form of independent particles. These viral particles, also known as virions, consist of two or three parts: (i) the genetic material (genome) made from either DNA or RNA, long molecules that carry genetic information; (ii) a protein coat called the capsid, which surrounds and protects the genetic material; and in some cases (iii) an envelope of lipids that surrounds the protein coat when they are outside a cell. The shapes of these virus particles range from simple helical and icosahedral forms for some virus species to more complex structures for others. Most virus species have virions that are too small to be seen with an optical microscope. The average virion is about one one-hundredth the size of the average bacterium.

Background

Bacteriophages (phages)
Multiple phages attached to a bacterial cell wall at 200,000x magnification
Diagram of a typical tailed phage
                  Phage injecting its genome into bacteria

Viruses are found wherever there is life and have probably existed since living cells first evolved.[3] The origins of viruses in the evolutionary history of life are unclear because they do not form fossils. molecular techniques have been used to compare the DNA or RNA of viruses and are a useful means of investigating how they arose.[4] Some viruses may have evolved from plasmids—pieces of DNA that can move between cells—while others may have evolved from bacteria. In evolution, viruses are an important means of horizontal gene transfer, which increases genetic diversity.[5]

Viruses are now recognised as ancient and as having origins that pre-date the divergence of life into the three domains.[6]

Opinions differ on whether viruses are a form of life or organic structures that interact with living organisms.[7] They are considered by some to be a life form, because they carry genetic material, reproduce by creating multiple copies of themselves through self-assembly, and evolve through natural selection. However they lack key characteristics such as a cellular structure generally considered necessary to count as life. Because they possess some but not all such qualities, viruses have been described as replicators[8] and as "organisms at the edge of life".[9]

Marine phages

These are cyanophages, viruses that infect cyanobacteria (scale bars indicate 100 nm)
Further information: Marine bacteriophage

Bacteriophages, often just called phages, are viruses that parasite bacteria and archaea. Marine phages parasite marine bacteria and archaea, such as cyanobacteria.[10] They are a common and diverse group of viruses and are the most abundant biological entity in marine environments, because their hosts, bacteria, are typically the numerically dominant cellular life in the sea. Generally there are about 1 million to 10 million viruses in each mL of seawater, or about ten times more double-stranded DNA viruses than there are cellular organisms,[11][12] although estimates of viral abundance in seawater can vary over a wide range.[13][14] For a long time, tailed phages of the order Caudovirales seemed to dominate marine ecosystems in number and diversity of organisms.[10]

Virions of different families of tailed phages

However, as a result of more resent research, non-tailed viruses appear to be dominant in multiple depths and oceanic regions, followed by the Caudovirales families of myoviruses, podoviruses, and siphoviruses.[15] Phages belonging to the families Corticoviridae,[16] Inoviridae,[17] Microviridae,[18] and Autolykiviridae[19][20][21][22] are also known to infect diverse marine bacteria.

There are also archaean viruses which replicate within archaea: these are double-stranded DNA viruses with unusual and sometimes unique shapes.[23][24] These viruses have been studied in most detail in the thermophilic archaea, particularly the orders Sulfolobales and Thermoproteales.[25]

Role of marine viruses

Connections between the different compartments of the living (bacteria/viruses and phytoplankton/zooplankton) and the nonliving (DOM/POM and inorganic matter) environment [26]

Microorganisms make up about 70% of the marine biomass.[27] It is estimated viruses kill 20% of this biomass each day and that there are 15 times as many viruses in the oceans as there are bacteria and archaea. Viruses are the main agents responsible for the rapid destruction of harmful algal blooms,[28] which often kill other marine life.[29] The number of viruses in the oceans decreases further offshore and deeper into the water, where there are fewer host organisms.[30]

Viruses are an important natural means of transferring genes between different species, which increases genetic diversity and drives evolution.[5] It is thought that viruses played a central role in the early evolution, before the diversification of bacteria, archaea and eukaryotes, at the time of the last universal common ancestor of life on Earth.[31] Viruses are still one of the largest reservoirs of unexplored genetic diversity on Earth.[30]

Viruses are part of the hydrothermal vent microbial community and their influence on the microbial ecology in these ecosystems is a burgeoning field of research.[32] Viruses are the most abundant life in the ocean, harboring the greatest reservoir of genetic diversity.[33] As their infections are often fatal, they constitute a significant source of mortality and thus have widespread influence on biological oceanographic processes, evolution and biogeochemical cycling within the ocean.[34] Evidence has been found however to indicate that viruses found in vent habitats have adopted a more mutualistic than parasitic evolutionary strategy in order to survive the extreme and volatile environment they exist in.[35] Deep-sea hydrothermal vents were found to have high numbers of viruses indicating high viral production.[36] Like in other marine environments, deep-sea hydrothermal viruses affect abundance and diversity of prokaryotes and therefore impact microbial biogeochemical cycling by lysing their hosts to replicate.[37] However, in contrast to their role as a source of mortality and population control, viruses have also been postulated to enhance survival of prokaryotes in extreme environments, acting as reservoirs of genetic information. The interactions of the virosphere with microorganisms under environmental stresses is therefore thought to aide microorganism survival through dispersal of host genes through horizontal gene transfer.Cite error: A <ref> tag is missing the closing </ref> (see the help page). The DOM recycled by the viral shunt pathway is comparable to the amount generated by the other main sources of marine DOM.[38]

Viruses can easily infect microorganisms in the microbial loop due to their relative abundance compared to microbes.[39] Prokaryotic and eukaryotic mortality contribute to carbon nutrient recycling through cell lysis. There is evidence as well of nitrogen (specifically ammonium) regeneration. This nutrient recycling helps stimulates microbial growth.[40] As much as 25% of the primary production from phytoplankton in the global oceans may be recycled within the microbial loop through viral shunting.[41]

Giant marine viruses

The giant mimivirus
Largest known virus, Tupanvirus, named after Tupã, the Guarani supreme god of creation

Viruses normally range in length from about 20 to 300 nanometers. This can be contrasted with the length of bacteria, which starts at about 400 nanometers. There are also giant viruses, often called giruses, typically about 1000 nanometers (one micron) in length. All giant viruses belongto phylum Nucleocytoviricota (NCLDV), together with poxviruses. The largest known of these is Tupanvirus. This genus of giant virus was discovered in 2018 in the deep ocean as well as a soda lake, and can reach up to 2.3 microns in total length.[42]

The discovery and subsequent characterization of giant viruses has triggered some debate concerning their evolutionary origins. The two main hypotheses for their origin are that either they evolved from small viruses, picking up DNA from host organisms, or that they evolved from very complicated organisms into the current form which is not self-sufficient for reproduction.[43] What sort of complicated organism giant viruses might have diverged from is also a topic of debate. One proposal is that the origin point actually represents a fourth domain of life,[44][45] but this has been largely discounted.[46][47]

References

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