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[[File:Origin of eukaryotic viruses.webp|thumb|upright=1.5|left| The second melting pot of virus evolution: origin of eukaryotic viruses. Characteristic images of archaeal, bacterial, and eukaryotic viruses are shown.<ref name=Koonin2006 />]]
[[File:Origin of eukaryotic viruses.webp|thumb|upright=1.5|left| The second melting pot of virus evolution: origin of eukaryotic viruses. Characteristic images of archaeal, bacterial, and eukaryotic viruses are shown.<ref name=Koonin2006 />]]


; Marine protists
By 2015, about 40 viruses affecting [[marine protists]] had been isolated and examined, most of them viruses of microalgae.<ref>Tomaru Y, Kimura K and Nagasaki K (2015) [https://books.google.co.nz/books?id=vgyhCgAAQBAJ&pg=PA502&dq=%22Marine+Protist+Viruses%22&hl=en&newbks=1&newbks_redir=0&sa=X&ved=0ahUKEwjHksvVmurpAhWXIbcAHd3pDpEQ6AEIKDAA#v=onepage&q=%22Marine%20Protist%20Viruses%22&f=false "Marine Protist Viruses"]. In: Ohtsuka S, Suzaki T, Horiguchi T, Suzuki N, Not F (eds) ''Marine Protists'' pages 501–517. Springer, Tokyo. {{doi|10.1007/978-4-431-55130-0_20}}. {{ISBN|978-4-431-55130-0}}.</ref> The genomes of these marine protist viruses are highly diverse.<ref>Hyman, P. and Abedon, S.T. (2012) "Smaller fleas: viruses of microorganisms". ''Scientifica'', . {{doi|10.6064/2012/734023}}. [[File:CC-BY icon.svg|50px]] Material was copied from this source, which is available under a [https://creativecommons.org/licenses/by/3.0/ Creative Commons Attribution 3.0 International License].
</ref><ref>Short, S.M. (2012) "The ecology of viruses that infect eukaryotic algae". ''Environmental microbiology'', '''14'''(9): 2253–2271. {{doi|10.1111/j.1462-2920.2012.02706.x}}.</ref>

; Marine invertebrates
[[Marine invertebrates]] are suscepible to viral diseases.<ref>TJohnson, P.T. (1984) "Viral diseases of marine invertebrates". ''Helgoländer Meeresuntersuchungen'', '''37'''(1-4): 65–98. {{doi|10.1007/BF01989296}}.</ref><ref>Renault T (2011) [https://books.google.com/books?hl=en&lr=&id=8uXDYcIf1fwC&oi=fnd&pg=PA153&dq=Viruses+infecting+marine+molluscs&ots=ewtZFuDD7X&sig=_LUTUTSCkU0L4ugAw4ukdUpip8A#v=onepage&q=Viruses%20infecting%20marine%20molluscs&f=false "Viruses infecting marine molluscs"] In: Hurst CJ (Ed) ''Studies in Viral Ecology'', Volume 2: Animal Host Systems, John Wiley & Sons. {{ISBN|9781118024584}}.</ref><ref>Arzul, I., Corbeil, S., Morga, B. and Renault, T., 2017. Viruses infecting marine molluscs. Journal of invertebrate pathology, 147, pp.118-135. {{doi|10.1016/j.jip.2017.01.009}}.</ref> [[Sea star wasting disease]] is a disease of [[starfish]] and several other echinoderms that appears sporadically, causing mass mortality of those affected.<ref>Dawsoni, Solaster. "Sea Star Species Affected by Wasting Syndrome." Pacificrockyintertidal.org Seastarwasting.org (n.d.): n. pag. ''Ecology and Evolutionary Biology''. Web.</ref> There are around 40 different species of sea stars that have been affected by this disease. In 2014 it was suggested that the disease is associated with a single-stranded DNA virus now known as the [[sea star-associated densovirus]] (SSaDV); however, sea star wasting disease is not fully understood.<ref name=MARINe2018>{{Cite web|url=https://www.eeb.ucsc.edu/pacificrockyintertidal/data-products/sea-star-wasting/|title=Sea Star Wasting Syndrome {{!}} MARINe|website=eeb.ucsc.edu|language=en|access-date=2018-06-03}}</ref>
[[Marine invertebrates]] are suscepible to viral diseases.<ref>TJohnson, P.T. (1984) "Viral diseases of marine invertebrates". ''Helgoländer Meeresuntersuchungen'', '''37'''(1-4): 65–98. {{doi|10.1007/BF01989296}}.</ref><ref>Renault T (2011) [https://books.google.com/books?hl=en&lr=&id=8uXDYcIf1fwC&oi=fnd&pg=PA153&dq=Viruses+infecting+marine+molluscs&ots=ewtZFuDD7X&sig=_LUTUTSCkU0L4ugAw4ukdUpip8A#v=onepage&q=Viruses%20infecting%20marine%20molluscs&f=false "Viruses infecting marine molluscs"] In: Hurst CJ (Ed) ''Studies in Viral Ecology'', Volume 2: Animal Host Systems, John Wiley & Sons. {{ISBN|9781118024584}}.</ref><ref>Arzul, I., Corbeil, S., Morga, B. and Renault, T., 2017. Viruses infecting marine molluscs. Journal of invertebrate pathology, 147, pp.118-135. {{doi|10.1016/j.jip.2017.01.009}}.</ref> [[Sea star wasting disease]] is a disease of [[starfish]] and several other echinoderms that appears sporadically, causing mass mortality of those affected.<ref>Dawsoni, Solaster. "Sea Star Species Affected by Wasting Syndrome." Pacificrockyintertidal.org Seastarwasting.org (n.d.): n. pag. ''Ecology and Evolutionary Biology''. Web.</ref> There are around 40 different species of sea stars that have been affected by this disease. In 2014 it was suggested that the disease is associated with a single-stranded DNA virus now known as the [[sea star-associated densovirus]] (SSaDV); however, sea star wasting disease is not fully understood.<ref name=MARINe2018>{{Cite web|url=https://www.eeb.ucsc.edu/pacificrockyintertidal/data-products/sea-star-wasting/|title=Sea Star Wasting Syndrome {{!}} MARINe|website=eeb.ucsc.edu|language=en|access-date=2018-06-03}}</ref>


; Marine vertebrates
[[Fish]] are particularly prone to infections with [[rhabdovirus]]es, which are distinct from, but related to rabies virus. At least nine types of rhabdovirus cause economically important diseases in species including salmon, pike, perch, sea bass, carp and cod. The symptoms include anaemia, bleeding, lethargy and a mortality rate that is affected by the temperature of the water. In hatcheries the diseases are often controlled by increasing the temperature to 15–18&nbsp;°C.<ref name=Murphy1999>{{cite book |author1=Murphy, FA |author2=Gibbs, EPJ |author3=Horzinek, MC |author4=Studdart MJ |title=Veterinary Virology |publisher=Academic Press |location=Boston |year=1999 |isbn=978-0-12-511340-3}}</ref>{{rp|442–443}} Like all vertebrates, fish suffer from [[Herpesviridae|herpes viruses]]. These ancient viruses have co-evolved with their hosts and are highly species-specific.<ref name=Murphy1999 />{{rp|324}} In fish, they cause cancerous tumours and non-cancerous growths called [[hyperplasia]].<ref name=Murphy1999 />{{rp|325}}
[[Fish]] are particularly prone to infections with [[rhabdovirus]]es, which are distinct from, but related to rabies virus. At least nine types of rhabdovirus cause economically important diseases in species including salmon, pike, perch, sea bass, carp and cod. The symptoms include anaemia, bleeding, lethargy and a mortality rate that is affected by the temperature of the water. In hatcheries the diseases are often controlled by increasing the temperature to 15–18&nbsp;°C.<ref name=Murphy1999>{{cite book |author1=Murphy, FA |author2=Gibbs, EPJ |author3=Horzinek, MC |author4=Studdart MJ |title=Veterinary Virology |publisher=Academic Press |location=Boston |year=1999 |isbn=978-0-12-511340-3}}</ref>{{rp|442–443}} Like all vertebrates, fish suffer from [[Herpesviridae|herpes viruses]]. These ancient viruses have co-evolved with their hosts and are highly species-specific.<ref name=Murphy1999 />{{rp|324}} In fish, they cause cancerous tumours and non-cancerous growths called [[hyperplasia]].<ref name=Murphy1999 />{{rp|325}}


Revision as of 08:27, 5 June 2020

Structure of a typical virus, in this case a phage.[1] The appearance of these viruses has been likened to a miniature lunar lander.[2] They are essential to the regulation of marine ecosystems.[3]
Part of a series of overviews on
Marine life

Marine viruses are defined by their habitat as viruses that live in marine environments, that is, in the saltwater of seas or oceans or the brackish water of coastal estuaries. A virus is a small infectious agent that replicates only inside the living cells of another organism. Viruses can infect all types of life forms, from animals and plants to microorganisms, including bacteria and archaea as well as other viruses.[4]

When not inside a cell or in the process of infecting a cell, viruses exist in the form of independent particles. A single viral particle is called a virion. A viron 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. 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 linear size of the average bacterium.

A teaspoon of seawater typically contains about ten million marine viruses. Most of these viruses are bacteriophages which infect and destroy marine bacteria and control the growth of phytoplankton at the base of the marine food web. Bacteriophages are harmless to plants and animals, but are essential to the regulation of marine ecosystems. They supply key mechanisms for recycling ocean carbon and nutrients. In a process known as the viral shunt, organic molecules released from dead bacterial cells stimulate fresh bacterial and algal growth. In particular the breaking down of bacteria by viruses (lysis) has been shown to enhance nitrogen cycling and stimulate phytoplankton growth. Viral activity also affects the biological pump, the process which sequesters carbon in the deep ocean. By increasing the amount of respiration in the oceans, viruses are indirectly responsible for reducing the amount of carbon dioxide in the atmosphere by approximately 3 gigatonnes of carbon per year

Marine microorganisms contribute about 70% of the total marine biomass. It is estimated marine viruses kill 20% of this biomass every day. Viruses are the main agents responsible for the rapid destruction of harmful algal blooms which often kill other marine life. The number of viruses in the oceans decreases further offshore and deeper into the water, where there are fewer host organisms. Viruses are an important natural means of transferring genes between different species, which increases genetic diversity and drives evolution. It is thought viruses played a central role in early evolution before the diversification of bacteria, archaea and eukaryotes, at the time of the last universal common ancestor of life on Earth. Viruses are still one of the largest areas of unexplored genetic diversity on Earth.

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
Further information: introduction to viruses and viral evolution

Viruses are now recognised as ancient and as having origins that pre-date the divergence of life into the three domains.[5] They are found wherever there is life and have probably existed since living cells first evolved.[6] The origins of viruses in the evolutionary history of life are unclear because they do not form fossils. Molecular techniques are used to compare the DNA or RNA of viruses and are a useful means of investigating how they arose.[7] 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.[8]

Opinions differ on whether viruses are a form of life or organic structures that interact with living organisms.[9] 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[10] and as "organisms at the edge of life".[11]

Bacteriophages

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

Bacteriophages, often just called phages, are viruses that parasite bacteria. Marine phages parasite marine bacteria such as cyanobacteria.[12] They are a diverse group of viruses which are the most abundant biological entity in marine environments, because their hosts, bacteria, are typically the numerically dominant cellular life in the sea. There are up to ten times more phages in the oceans than there are bacteria,[13] reaching levels of 250 million bacteriophages per millilitre of seawater.[14] These viruses infect specific bacteria by binding to surface receptor molecules and then entering the cell. Within a short amount of time, in some cases just minutes, bacterial polymerase starts translating viral mRNA into protein. These proteins go on to become either new virions within the cell, helper proteins, which help assembly of new virions, or proteins involved in cell lysis. Viral enzymes aid in the breakdown of the cell membrane, and there are phages that can replicate three hundred phages twenty minutes after injection.[15]

Bacteria defend themselves from bacteriophages is by producing enzymes that destroy foreign DNA. These enzymes, called restriction endonucleases, cut up the viral DNA that bacteriophages inject into bacterial cells.[16] Bacteria also contain a system that uses CRISPR sequences to retain fragments of the genomes of viruses that the bacteria have come into contact with in the past, which allows them to block the virus's replication through a form of RNA interference.[17][18] This genetic system provides bacteria with acquired immunity to infection.[19]

Adsorption of cyanophages onto a marine Prochlorococcus
(a) Slice (~20 nm) through a reconstructed tomogram of P-SSP7 phage incubated with MED4, imaged at ~86 min post-infection. FC and EC show full-DNA capsid phage and empty capsid phage, respectively.
(b) same image visualised by highlighting the cell wall in orange, the plasma membrane in light yellow, the thylakoid membrane in green, carboxysomes in cyan, the polyphosphate body in blue, adsorbed phages on the sides or top of the cell in red, and cytoplasmic granules (probably mostly ribosomes) in light purple.[20]
scale bar: 200 nm
Process of a phage "landing" on a bacterium
The phage first adheres to the cell surface with its tail parallel to or leaning at an angle to the cell surface in the pre-infection stage. The tail then firmly stands on the cell surface and extends its fibers horizontally, rendering the phage infection-competent, after which viral DNA is released into the cell through an extensible tube.[20]
based on observations of the model cyanophage P-SSP7
interacting with the marine Prochlorococcus MED4 bacterium
Virions of different families of tailed phages

Microbes drive the nutrient transformations that sustain Earth’s ecosystems,[21] and the viruses that infect these microbes modulate both microbial population size and diversity.[22][20] The cyanobacterium Prochlorococcus, the most abundant oxygenic phototroph on Earth, contributes a substantial fraction of global primary carbon production, and often reaches densities of over 100,000 cells per milliliter in oligotrophic and temperate oceans.[23] Hence, viral (cyanophage) infection and lysis of Prochlorococcus represent an important component of the global carbon cycle. In addition to their ecological role in inducing host mortality, cyanophages influence the metabolism and evolution of their hosts by co-opting and exchanging genes, including core photosynthesis genes.[20]

For a long time, tailed phages of the order Caudovirales seemed to dominate marine ecosystems in number and diversity of organisms.[12] However, as a result of more recent research, non-tailed viruses appear to be dominant in multiple depths and oceanic regions, followed by the Caudovirales families of myoviruses, podoviruses, and siphoviruses.[24] Phages belonging to the families Corticoviridae,[25] Inoviridae,[26] Microviridae,[27] and Autolykiviridae[28][29][30][31] are also known to infect diverse marine bacteria.

Archaeal viruses

Evolution of the virus world: origin of the main lineages from the primordial gene pool
Characteristic images of RNA and protein structures are shown for each postulated stage of evolution, and characteristic virion images are shown for the emerging classes of viruses. Thin arrows show the postulated movement of genetic pools between inorganic compartments. Block arrows show the origin of different classes of viruses at different stages of pre-cellular evolution.[4]
Further information: archaeal viruses

Archaean viruses replicate within archaea: these are double-stranded DNA viruses with unusual and sometimes unique shapes.[32][33] These viruses have been studied in most detail in the thermophilic archaea, particularly the orders Sulfolobales and Thermoproteales.[34] Defences against these viruses involve RNA interference from repetitive DNA sequences within archaean genomes that are related to the genes of the viruses.[35][36] Most archaea have CRISPR–Cas systems as an adaptive defence against viruses. These enable archaea to retain sections of viral DNA, which are then used to target and eliminate subsequent infections by the virus using a process similar to RNA interference.[37]

Fungal viruses

Further information: mycovirus

Mycoviruses, also known as mycophages, are viruses that infect fungi. The infection of fungal cells is different from that of animal cells. Fungi have a rigid cell wall made of chitin, so most viruses can get inside these cells only after trauma to the cell wall.[38]

Eukaryote viruses

The second melting pot of virus evolution: origin of eukaryotic viruses. Characteristic images of archaeal, bacterial, and eukaryotic viruses are shown.[4]
Marine protists

By 2015, about 40 viruses affecting marine protists had been isolated and examined, most of them viruses of microalgae.[39] The genomes of these marine protist viruses are highly diverse.[40][41]

Marine invertebrates

Marine invertebrates are suscepible to viral diseases.[42][43][44] Sea star wasting disease is a disease of starfish and several other echinoderms that appears sporadically, causing mass mortality of those affected.[45] There are around 40 different species of sea stars that have been affected by this disease. In 2014 it was suggested that the disease is associated with a single-stranded DNA virus now known as the sea star-associated densovirus (SSaDV); however, sea star wasting disease is not fully understood.[46]

Marine vertebrates

Fish are particularly prone to infections with rhabdoviruses, which are distinct from, but related to rabies virus. At least nine types of rhabdovirus cause economically important diseases in species including salmon, pike, perch, sea bass, carp and cod. The symptoms include anaemia, bleeding, lethargy and a mortality rate that is affected by the temperature of the water. In hatcheries the diseases are often controlled by increasing the temperature to 15–18 °C.[47]: 442–443  Like all vertebrates, fish suffer from herpes viruses. These ancient viruses have co-evolved with their hosts and are highly species-specific.[47]: 324  In fish, they cause cancerous tumours and non-cancerous growths called hyperplasia.[47]: 325 

In 1984, infectious salmon anemia (ISAv) was discovered in Norway in an Atlantic salmon hatchery. Eighty percent of the fish in the outbreak died. ISAv, a viral disease, is now a major threat to the viability of Atlantic salmon farming.[48] As the name implies, it causes severe anemia of infected fish. Unlike mammals, the red blood cells of fish have DNA, and can become infected with viruses. Management strategies include developing a vaccine and improving genetic resistance to the disease.[49]

Marine mammals are also susceptible to marine viral infections. In 1988 and 2002, thousands of harbour seals were killed in Europe by phocine distemper virus.[50] Many other viruses, including caliciviruses, herpesviruses, adenoviruses and parvoviruses, circulate in marine mammal populations.[51]

Giant marine viruses

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

Most viruses 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.[52]

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.[53] 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,[54][55] but this has been largely discounted.[56][57]

Virophages

Further information: Virophages

Virophages are small, double-stranded DNA viral phages that require the co-infection of another virus. The co-infecting viruses are typically giant viruses. Virophages rely on the viral replication factory of the co-infecting giant virus for their own replication. One of the characteristics of virophages is that they have a parasitic relationship with the co-infecting virus. Their dependence upon the giant virus for replication often results in the deactivation of the giant viruses. The virophage may improve the recovery and survival of the host organism. Unlike satellite viruses, virophages have a parasitic effect on their co-infecting virus. Virophages have been observed to render a giant virus inactive and thereby improve the condition of the host organism.

All known virophages are grouped into the family Lavidaviridae (from "large virus dependent or associated" + -viridae).[59] The first virophage was discovered in a cooling tower in Paris, France in 2008. It was discovered with its co-infecting giant virus, Acanthamoeba castellanii mamavirus (ACMV). The virophage was named Sputnik and its replication relied entirely on the co-infection of ACMV and its cytoplasmic replication machinery. Sputnik was also discovered to have an inhibitory effect on ACMV and improved the survival of the host. Other characterised virophages include Sputnik 2, Sputnik 3, Zamilon and Mavirus.[60][61]

A majority of these virophages are being discovered by analyzing metagenomic data sets. In metagenomic analysis, DNA sequences are run through multiple bioinformatic algorithms which pull out certain important patterns and characteristics. In these data sets are giant viruses and virophages. They are separated by looking for sequences around 17 to 20 kbp long which have similarities to already sequenced virophages. These virophages can have linear or circular double-stranded DNA genomes.[62] Virophages in culture have icosahedral capsid particles that measure around 40 to 80 nanometers long.[63] Virophage particles are so small that electron microscopy must be used to view these particles. Metagenomic sequence-based analyses have been used to predict around 57 complete and partial virophage genomes[64] and in December 2019 to identify 328 high-quality (complete or near-complete) genomes from diverse habitats including the human gut, plant rhizosphere, and terrestrial subsurface, from 27 distinct taxonomic clades.[65]

The giant virus CroV attacks C.roenbergensis
Cafeteria roenbergensis a bacterivorous marine flagellate
A Mavirus virophage (lower left) lurking alongside a giant CroV [66]

A giant marine virus CroV infects and causes the death by lysis of the marine zooflagellate Cafeteria roenbergensis.[67] This impacts coastal ecology because Cafeteria roenbergensis feeds on bacteria found in the water. When there are low numbers of Cafeteria roenbergensis due to extensive CroV infections, the bacterial populations rise exponentially.[68] The impact of CroV on natural populations of C. roenbergensis remains unknown; however, the virus has been found to be very host specific, and does not infect other closely related organisms.[69] Cafeteria roenbergensis is also infected by a second virus, the Mavirus virophage, which is a satellite virus, meaning it is able to replicate only in the presence of another specific virus, in this case in the presence of CroV.[60] This virus interferes with the replication of CroV, which leads to the survival of C. roenbergensis cells. Mavirus is able to integrate into the genome of cells of C. roenbergensis, and thereby confer immunity to the population.[61]

Role of marine viruses

Virus-host interactions in the marine ecosystem,
including viral infection of bacteria, phytoplankton and fish[70]
The cycling of marine phytoplankton is helped by viral lysis [71]

Viruses are the most abundant biological entity in aquatic environments[4] On average there are about ten million of them in a typical teaspoon of seawater.[72] Most of these viruses are bacteriophages infecting heterotrophic bacteria and cyanophages infecting cyanobacteria. They are essential to the regulation of marine ecosystems.[3]

Bacteriophages are harmless to plants and animals, and are essential to the regulation of marine and freshwater ecosystems[73] are important mortality agents of phytoplankton, the base of the foodchain in aquatic environments.[74] They infect and destroy bacteria in aquatic microbial communities, and are one of the most important mechanisms of recycling carbon and nutrient cycling in marine environments. The organic molecules released from the dead bacterial cells stimulate fresh bacterial and algal growth, in a process known as the viral shunt.[75] In particular, lysis of bacteria by viruses has been shown to enhance nitrogen cycling and stimulate phytoplankton growth.[76]

Viral activity also affects the biological pump, the process where carbon is sequestered in the deep ocean.[51] By increasing the amount of respiration in the oceans, viruses are indirectly responsible for reducing the amount of carbon dioxide in the atmosphere by about three gigatonnes of carbon per year.[51]

Microorganisms make up about 70% of the marine biomass.[77] 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,[78] which often kill other marine life.[79] The number of viruses in the oceans decreases further offshore and deeper into the water, where there are fewer host organisms.[80]

Viruses are an important natural means of transferring genes between different species, which increases genetic diversity and drives evolution.[8] It is thought viruses played a central role in early evolution, before bacteria, archaea and eukaryotes diversified, at the time of the last universal common ancestor of life on Earth.[81] Viruses are still one of the largest reservoirs of unexplored genetic diversity on Earth.[80]

Connections between the different compartments of the living (bacteria/viruses and phytoplankton/zooplankton) and the nonliving (DOM/POM and inorganic matter) environment [82]

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.[83] Viruses are the most abundant life in the ocean, harboring the greatest reservoir of genetic diversity.[78] 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.[80] 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.[84] Deep-sea hydrothermal vents were found to have high numbers of viruses indicating high viral production.[85] 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.[86] 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.[87]

Viral shunt

The viral shunt pathway facilitates the flow of dissolved organic matter (DOM) and particulate organic matter (POM) through the marine food web
Main article: Viral shunt

The viral shunt pathway is a mechanism that prevents (prokaryotic and eukaryotic) marine microbial particulate organic matter (POM) from migrating up trophic levels by recycling them into dissolved organic matter (DOM), which can be readily taken up by microorganisms. Viral shunting helps maintain diversity within the microbial ecosystem by preventing a single species of marine microbe from dominating the micro-environment.[88] The DOM recycled by the viral shunt pathway is comparable to the amount generated by the other main sources of marine DOM.[89]

Viruses can easily infect microorganisms in the microbial loop due to their relative abundance compared to microbes.[90] 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.[91] As much as 25% of the primary production from phytoplankton in the global oceans may be recycled within the microbial loop through viral shunting.[92]

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