Structure of a typical virus, in this case aphage.[1] The appearance of these viruses has been likened to a miniature lunar lander.[2] Marine viruses are essential to the regulation of marine ecosystems.[3]
When not inside a cell or in the process of infecting a cell, viruses exist in the form of independent particles calledvirions. A virion contains agenome (a longmolecule that carries genetic information in the form of eitherDNA orRNA) surrounded by acapsid (aprotein coat protecting the genetic material). The shapes of these virus particles range from simplehelical andicosahedral forms for some virus species to more complex structures for others. Most virus species have virions that are too small to be seen with anoptical microscope. The average virion is about one one-hundredth the linear size of the averagebacterium.
A teaspoon of seawater typically contains about fifty million viruses.[6] Most of these viruses arebacteriophages which infect and destroymarine bacteria and control the growth ofphytoplankton at the base of themarine food web. Bacteriophages are harmless to plants and animals but are essential to the regulation of marine ecosystems. They supply key mechanisms forrecycling ocean carbon andnutrients. In a process known as theviral 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 enhancenitrogen cycling and stimulate phytoplankton growth. Viral activity also affects thebiological pump, the process whichsequesters 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 3gigatonnes of carbon per year.
Marine microorganisms make up about 70% of the totalmarine biomass. It is estimated marine viruses kill 20% of the microorganism biomass every day. Viruses are the main agents responsible for the rapid destruction of harmfulalgal blooms which often kill othermarine 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 oftransferring genes between different species, which increasesgenetic 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.
Viruses are now recognised as ancient and as having origins that pre-date the divergence of life into thethree domains.[7] They are found wherever there is life and have probably existed since living cells first evolved.[8] The origins of viruses in theevolutionary 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.[9] Some viruses may haveevolved fromplasmids—pieces of DNA that can move between cells—while others may have evolved from bacteria. In evolution, viruses are an important means ofhorizontal gene transfer, which increasesgenetic diversity.[10]
Opinions differ on whether viruses are a form oflife or organic structures that interact with living organisms.[11] 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 throughnatural 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[12] and as "organisms at the edge of life".[13]
The existence of viruses in the ocean was discovered throughelectron microscopy andepifluorescence microscopy of ecological water samples, and later throughmetagenomic sampling of uncultured viral samples.[14][15] Marine viruses, although microscopic and essentially unnoticed byscientists until recently, are the most abundant and diverse biological entities in the ocean. Viruses have an estimated abundance of 1030 in the ocean, or between 106 and 1011 viruses permillilitre.[4] Quantification of marine viruses was originally performed using transmission electron microscopy but has been replaced by epifluorescence orflow cytometry.[16]
Bacteriophages, often contracted tophages, are viruses thatparasitize bacteria for replication. As aptly named, marine phages parasitizemarine bacteria, such ascyanobacteria.[17] 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,[18] reaching levels of 250 million bacteriophages per millilitre of seawater.[19] These viruses infect specific bacteria by binding tosurface receptor molecules and then entering the cell. Within a short amount of time, in some cases just minutes, bacterialpolymerase 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.[20]
Bacteriophages (phages)
Multiple phages attached to a bacterial cell wall at 200,000x magnification
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.[21]
based on observations of the model cyanophage P-SSP7 interacting with the marineProchlorococcus MED4 bacterium
(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.[21]
scale bar: 200 nm
Bacteria defend themselves from bacteriophages by producing enzymes that destroy foreign DNA. These enzymes, calledrestriction endonucleases, cut up the viral DNA that bacteriophages inject into bacterial cells.[22] Bacteria also contain a system that usesCRISPR 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 ofRNA interference.[23][24] This genetic system provides bacteria withacquired immunity to infection.[25]
Thelytic cycle, the reproductive cycle of the bacteriophage, has six stages: → attachment: the phage attaches itself to the surface of the host cell → penetration: the phage injects its DNA through the cell membrane → transcription: the host cell's DNA is degraded and the cell's metabolism is directed to initiate phage biosynthesis → biosynthesis: the phage DNA replicates inside the cell → maturation: the replicated material assembles into fully formed viral phages →lysis: the newly formed phages are released from the infected cell (which is itself destroyed in the process) to seek out new host cells[26]
Microbes drive the nutrient transformations that sustain Earth's ecosystems,[27] and the viruses that infect these microbes modulate both microbial population size and diversity.[28][21] ThecyanobacteriumProchlorococcus, the most abundant oxygenicphototroph on Earth, contributes a substantial fraction of global primary carbon production, and often reaches densities of over 100,000 cells per milliliter inoligotrophic and temperate oceans.[29] Hence, viral (cyanophage) infection and lysis ofProchlorococcus represent an important component of the globalcarbon 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.[21]
For a long time,tailed phages of the orderCaudovirales seemed to dominate marine ecosystems in number and diversity of organisms.[17] However, as a result of more recent research, non-tailed viruses appear to dominate multiple depths and oceanic regions.[31] These non-tailed phages also infect marine bacteria, and include the familiesCorticoviridae,[32]Inoviridae,[33]Microviridae[34] andAutolykiviridae.[35][36][37][38]
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.[5]
Archaean viruses replicate withinarchaea: these are double-stranded DNA viruses with unusual and sometimes unique shapes.[40][41] These viruses have been studied in most detail in thethermophilic archaea, particularly the ordersSulfolobales andThermoproteales.[42] Defences against these viruses involve RNA interference fromrepetitive DNA sequences within archaean genomes that are related to the genes of the viruses.[43][44] 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.[45]
Mycoviruses, also known as mycophages, are viruses that infectfungi. 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.[46]
See, Nerva L, Ciuffo M, Vallino M, Margaria P, Varese G, Gnavi G, Turina M (2016). "Multiple approaches for the detection and characterization of viral and plasmid symbionts from a collection of marine fungi".Virus Research.219:22–38.doi:10.1016/j.virusres.2015.10.028.hdl:2318/1527617.PMID26546154.S2CID53417720.
The second melting pot of virus evolution: origin of eukaryotic viruses. Characteristic images of archaeal, bacterial, and eukaryotic viruses are shown.[5]
By 2015, about 40 viruses affectingmarine protists had been isolated and examined, most of them viruses of microalgae.[47] The genomes of these marine protist viruses are highly diverse.[48][49]Marine algae can be infected by viruses in the familyPhycodnaviridae. These are large (100–560 kb)double-stranded DNA viruses withicosahedral shaped capsids. By 2014, 33 species divided into six genera had been identified within the family,[50][51] which belongs to a super-group of large viruses known asnucleocytoplasmic large DNA viruses. Evidence was published in 2014 suggesting some strains ofPhycodnaviridae might infect humans rather than just algal species, as was previously believed.[52] Most genera under this family enter the host cell by cell receptorendocytosis and replicate in the nucleus.
Phycodnaviridae play important ecological roles by regulating the growth and productivity of their algal hosts. Algal species suchHeterosigma akashiwo and the genusChrysochromulina can form dense blooms which can be damaging to fisheries, resulting in losses in the aquaculture industry.[53]Heterosigma akashiwo virus (HaV) has been suggested for use as a microbial agent to prevent the recurrence of toxic red tides produced by this algal species.[54] ThecoccolithovirusEmiliania huxleyi virus 86, a giant double-strandedDNA virus, infects the ubiquitouscoccolithophoreEmiliania huxleyi.[50][51] This virus has one of the largest known genomes among marine viruses.[55]Phycodnaviridae cause death and lysis of freshwater and marine algal species, liberating organic carbon, nitrogen and phosphorus into the water, providing nutrients for themicrobial loop.[56]
The virus-to-prokaryote ratio, VPR, is often used as an indicator of the relationship between viruses and hosts. Studies have used VPR to indirectly infer virus impact on marine microbial productivity, mortality, and biogeochemical cycling.[57] However, in making these approximations, scientists assume a VPR of 10:1, the median observed VPR in the surface ocean.[57][18] The actual VPR varies greatly depending on location, so VPR may not be the accurate proxy for viral activity or abundance as it has been treated.[57][58]
Virus-host interactions in the marine ecosystem, including viral infection of bacteria, phytoplankton and fish[59]
Marine invertebrates are susceptible to viral diseases.[60][61][62]Sea star wasting disease is a disease ofstarfish and several other echinoderms that appears sporadically, causing mass mortality of those affected.[63] 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 thesea star-associated densovirus (SSaDV); however, sea star wasting disease is not fully understood.[64]
Fish are particularly prone to infections withrhabdoviruses, 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 (59–64 °F).[65]: 442–443 Like all vertebrates, fish suffer fromherpes viruses. These ancient viruses have co-evolved with their hosts and are highly species-specific.[65]: 324 In fish, they cause cancerous tumours and non-cancerous growths calledhyperplasia.[65]: 325
In 1984,infectious salmon anemia (ISAv) was discovered in Norway in anAtlantic salmon hatchery. Eighty per cent of the fish in the outbreak died. ISAv, a viral disease, is now a major threat to the viability of Atlantic salmon farming.[66] As the name implies, it causes severeanemia of infectedfish. 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.[67]
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 alsogiant viruses, often calledgiruses, typically about 1000 nanometers (one micron) in length. All giant viruses belong to thephylumNucleocytoviricota (NCLDV), together withpoxviruses.The largest known of these isTupanvirus. 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.[70]
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.[72] 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 fourthdomain of life,[73][74] but this has been largely discounted.[75][76]
Virophages are small, double-stranded DNA viruses that rely on theco-infection ofgiant 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 aparasitic 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 othersatellite viruses, virophages have aparasitic 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 familyLavidaviridae (from "large virus dependent or associated" +-viridae).[77] The first virophage was discovered in acooling tower in Paris in 2008. It was discovered with its co-infecting giant virus,Acanthamoeba castellaniimamavirus (ACMV). The virophage was namedSputnik 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 andMavirus.[78][79]
Most of these virophages were discovered by analyzingmetagenomic 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.[80] Virophages in culture have icosahedral capsid particles that measure around 40 to 80 nanometers long.[81] 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[82] 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.[83]
A giant marine virusCroV infects and causes the death bylysis of the marine zooflagellateCafeteria roenbergensis.[85] This impacts coastal ecology becauseCafeteria roenbergensis feeds on bacteria found in the water. When there are low numbers ofCafeteria roenbergensis due to extensive CroV infections, the bacterial populations rise exponentially.[86] The impact ofCroV on natural populations ofC. roenbergensis remains unknown; however, the virus has been found to be very host specific, and does not infect other closely related organisms.[87] Cafeteria roenbergensis is also infected by a second virus, theMavirus virophage, during co-infection with CroV.[78] This virus interferes with the replication of CroV, which leads to the survival ofC. roenbergensis cells.Mavirus is able to integrate into the genome of cells ofC. roenbergensis and thereby confer immunity to the population.[79]
Although marine viruses have only recently been studied extensively, they are already known to hold critical roles in many ecosystem functions and cycles.[88] Marine viruses offer a number of importantecosystem services and are essential to the regulation of marine ecosystems.[3] Marine bacteriophages and other viruses appear to influencebiogeochemical cycles globally, provide and regulatemicrobial biodiversity,cycle carbon through marinefood webs, and are essential in preventing bacterialpopulation explosions.[89]
The dominant hosts for viruses in the ocean are marine microorganisms, such as bacteria.[14] Bacteriophages are harmless to plants and animals, and are essential to the regulation of marine and freshwater ecosystems[91] are important mortality agents ofphytoplankton, the base of thefoodchain in aquatic environments.[92] They infect and destroy bacteria in aquatic microbial communities, and are one of the most important mechanisms ofrecycling carbon andnutrient cycling in marine environments. The organic molecules released from the dead bacterial cells stimulate fresh bacterial and algal growth, in a process known as theviral shunt.[93]
In this way, marine viruses are thought to play an important role in nutrient cycles by increasing the efficiency of thebiological pump. Viruses causelysis of living cells, that is, they break the cell membranes down. This releases compounds such asamino acids andnucleic acids, which tend to be recycled near the surface.
Viral activity also enhances the ability of the biological pump tosequester carbon in the deep ocean.[69] Lysis releases more indigestible carbon-rich material like that found in cell walls, which is likely exported to deeper waters. Thus, the material that is exported to deeper waters by the viral shunt is probably more carbon rich than the material from which it was derived.[94][95] 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 threegigatonnes of carbon per year.[69] Lysis of bacteria by viruses has been shown to also enhance nitrogen cycling and stimulate phytoplankton growth.[96]
Theviral shunt pathway is a mechanism that prevents (prokaryotic andeukaryotic)marine microbialparticulate organic matter (POM) from migrating uptrophic levels by recycling them intodissolved 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.[97] The DOM recycled by the viral shunt pathway is comparable to the amount generated by the other main sources of marine DOM.[98]
Theviral shunt pathway facilitates the flow ofdissolved organic matter (DOM) andparticulate organic matter (POM) through the marine food web. Connections between the different compartments of the living (bacteria/viruses and phytoplankton/zooplankton) and the nonliving (DOM/POM and inorganic matter) environment[99]
Viruses are the most abundant biological entity in marine environments.[5] On average there are about ten million of them in one milliliter of seawater.[100] Most of these viruses arebacteriophages infecting heterotrophic bacteria andcyanophages infecting cyanobacteria. Viruses easily infect microorganisms in themicrobial loop due to their relative abundance compared to microbes.[101] Prokaryotic and eukaryotic mortality contribute to carbon nutrient recycling throughcell lysis. There is evidence as well of nitrogen (specifically ammonium) regeneration. This nutrient recycling helps stimulates microbial growth.[102] As much as 25% of the primary production from phytoplankton in the global oceans may be recycled within the microbial loop through viral shunting.[103]
Viruses act as "regulators" of the globalcarbon cycle because they impact the material cycles and energy flows offood webs and themicrobial loop. The average contribution of viruses to the Earth ecosystem carbon cycle is 8.6%, of which its contribution to marine ecosystems (1.4%) is less than its contribution to terrestrial (6.7%) and freshwater (17.8%) ecosystems. Over the past 2,000 years, anthropogenic activities andclimate change have gradually altered the regulatory role of viruses in ecosystem carbon cycling processes. This has been particularly conspicuous over the past 200 years due to rapid industrialization and the attendant population growth.[104]
Comparison of how virus regulate the carbon cycle in terrestrial ecosystems (left) and in marine ecosystems (right). Arrows show the roles viruses play in the traditional food web, the microbial loop and the carbon cycle. Light green arrows represent the traditional food web, white arrows represent the microbial loop, and white dotted arrows represent the contribution rate of carbon produced byviral lysing of bacteria to the ecosystemdissolved organic carbon (DOC) pool. Freshwater ecosystems are regulated in a manner similar to marine ecosystems, and are not shown separately. The microbial loop is an important supplement to the classic food chain, wherein dissolved organic matter is ingested byheterotrophic "planktonic" bacteria duringsecondary production. These bacteria are then consumed byprotozoa,copepods and other organisms, and eventually returned to the classical food chain.[104]
Microorganisms make up about 70% of the marine biomass.[105] It is estimated viruses kill 20% of the microorganism 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 harmfulalgal blooms,[106] which often kill other marine life.[107] Scientists are exploring the potential of marinecyanophages to be used to prevent or reverseeutrophication. The number of viruses in the oceans decreases further offshore and deeper into the water, where there are fewer host organisms.[108]
Marine bacteriophages often containauxiliary metabolic genes, host-derived genes thought to sustain viral replication by supplementing host metabolism during viral infection.[109] These genes can impact multiple biogeochemical cycles, including carbon, phosphorus, sulfur, and nitrogen.[110][111][112][113]
Viruses are an important natural means oftransferring genes between different species, which increasesgenetic diversity and drives evolution.[10] It is thought viruses played a central role in early evolution, before bacteria, archaea and eukaryotes diversified, at the time of thelast universal common ancestor of life on Earth.[114] Viruses are still one of the largest reservoirs of unexplored genetic diversity on Earth.[108]
Marine coastal habitats sit at the interface between the land and the ocean. It is likely thatRNA viruses play significant roles in these environments.[115]
Marine surface habitats sit at the interface between the atmosphere and the ocean. Thebiofilm-like habitat at the surface of the ocean harbours surface-dwelling microorganisms, commonly referred to asneuston. Viruses in the microlayer, the so-calledvirioneuston, have recently become of interest to researchers as enigmatic biological entities in the boundary surface layers with potentially important ecological impacts. Given this vast air–water interface sits at the intersection of major air–water exchange processes spanning more than 70% of the global surface area, it is likely to have profound implications formarine biogeochemical cycles, on themicrobial loop and gas exchange, as well as themarine food web structure, the global dispersal of airborne viruses originating from thesea surface microlayer, and human health.[116]
Marine viral activity presents a potential explanation of theparadox of the plankton proposed byGeorge Hutchinson in 1961.[117] The paradox of the plankton is that many plankton species have been identified in small regions in the ocean where limited resources should createcompetitive exclusion, limiting the number of coexisting species.[117] Marine viruses could play a role in this effect, as viral infection increases as potential contact with hosts increases.[4] Viruses could therefore control the populations of plankton species that grow too abundant, allowing a wide diversity of species to coexist.[4]
Marine bacteriophages play an important role indeep seaecosystems. There are between 5x1012 and 1x1013 phages per square metre in deep sea sediments and their abundance closely correlates with the number ofprokaryotes found in the sediments. They are responsible for the death of 80% of the prokaryotes found in the sediments, and almost all of these deaths are caused by celllysis (bursting). This allows nitrogen, carbon, and phosphorus from the living cells to be converted into dissolved organic matter and detritus, contributing to the high rate of nutrient turnover in deep sea sediments. Because of the importance of deep sea sediments in biogeochemical cycles, marine bacteriophages influence thecarbon,nitrogen andphosphorus cycles. More research needs to be done to more precisely elucidate these influences.[118]
Viruses are part of thehydrothermal vent microbial community and their influence on the microbial ecology in these ecosystems is a burgeoning field of research.[119] Viruses are the most abundant life in the ocean, harboring the greatest reservoir of genetic diversity.[106] As their infections are often fatal, they constitute a significant source of mortality and thus have widespread influence on biological oceanographic processes,evolution andbiogeochemical cycling within the ocean.[108] Evidence has been found however to indicate that viruses found in vent habitats have adopted a moremutualistic thanparasitic evolutionary strategy in order to survive the extreme and volatile environment they exist in.[120] Deep-sea hydrothermal vents were found to have high numbers of viruses indicating high viral production.[121] Like in other marine environments,deep-sea hydrothermal viruses affect abundance and diversity ofprokaryotes and therefore impact microbial biogeochemical cycling bylysing their hosts to replicate.[122] 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 throughhorizontal gene transfer.[123]
In addition to varied topographies and in spite of an extremely cold climate, the polar aquatic regions are teeming withmicrobial life. Even in sub-glacial regions, cellular life has adapted to these extreme environments where perhaps there are traces of early microbes on Earth. As grazing bymacrofauna is limited in most of these polar regions, viruses are being recognised for their role as important agents of mortality, thereby influencing thebiogeochemical cycling ofnutrients that, in turn, impact community dynamics at seasonal and spatial scales.[124] The polar regions are characterised by truncated food webs, and the role of viruses in ecosystem function is likely to be even greater than elsewhere in themarine food web, yet their diversity is still relatively under-explored, and the way in which they affect polar communities is not well understood,[125] particularly in nutrient cycling.[126][127][128][124]
Viruses are highly host specific.[129] A marine virus is more likely to infect cooccurring organisms, those that live in the same region the virus lives in.[130] Therefore,biogeography is an important factor in a virion's ability to infect.
Knowledge of this variation in viral populations across spatiotemporal and other environmental gradients is supported by viral morphology, as determined by transmission electron microscopy (TEM). Non-tailed viruses appear to be dominant in multiple depths and oceanic regions, followed by the Caudoviralesmyoviruses, podoviruses, and siphoviruses.[31] However, viruses belonging to familiesCorticoviridae,[131]Inoviridae[132] andMicroviridae[133] are also known to infect diverse marine bacteria.Metagenomic evidence suggests that microviruses (icosahedral ssDNA phages) are particularly prevalent in marine habitats.[133]
Metagenomic approaches to assess viral diversity are often limited by a lack of reference sequences, leaving many sequences unannotated.[134] However, viral contigs are generated through direct sequencing of a viral fraction, typically generated after 0.02-um filtration of a marine water sample, or through bioinformatics approaches to identify viral contigs or viral genomes from a microbial metagenome. Novel tools to identify putative viral contigs, such as VirSorter[135] and VirFinder,[136] allow for the assessment of patterns of viral abundance, host range, and functional content of marine bacteriophage.[137][138]
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![Cryo-electron micrograph of the CroV giant virus [71] scale bar=0.2 μm](/mt/?noimg=&dark=on&url=https%3a%2f%2fwikipedia.org%2fwiki%2fFile%3aCroV_TEM_(cropped).jpg)
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