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The adjective viral dates to Louis Pasteur was unable to find a causative agent for rabies and speculated about a pathogen too small to be detected using a microscope.
Thus, he could pass a solution containing bacteria through the filter and completely remove them. His experiments showed that crushed leaf extracts from infected tobacco plants remain infectious after filtration.
Ivanovsky suggested the infection might be caused by a toxin produced by bacteria, but did not pursue the idea.
Beijerinck maintained that viruses were liquid in nature, a theory later discredited by Wendell Stanley , who proved they were particulate.
He accurately diluted a suspension of these viruses and discovered that the highest dilutions lowest virus concentrations , rather than killing all the bacteria, formed discrete areas of dead organisms.
Counting these areas and multiplying by the dilution factor allowed him to calculate the number of viruses in the original suspension.
The study of phages provided insights into the switching on and off of genes , and a useful mechanism for introducing foreign genes into bacteria.
By the end of the 19th century, viruses were defined in terms of their infectivity , their ability to be filtered, and their requirement for living hosts.
Viruses had been grown only in plants and animals. In , Ross Granville Harrison invented a method for growing tissue in lymph , and, in , E.
Lambert used this method to grow vaccinia virus in fragments of guinea pig corneal tissue. Maitland grew vaccinia virus in suspensions of minced hens' kidneys.
Their method was not widely adopted until the s, when poliovirus was grown on a large scale for vaccine production. Another breakthrough came in , when the American pathologist Ernest William Goodpasture and Alice Miles Woodruff grew influenza and several other viruses in fertilised chickens' eggs.
This work enabled Jonas Salk to make an effective polio vaccine. The first images of viruses were obtained upon the invention of electron microscopy in by the German engineers Ernst Ruska and Max Knoll.
The first X-ray diffraction pictures of the crystallised virus were obtained by Bernal and Fankuchen in On the basis of her pictures, Rosalind Franklin discovered the full structure of the virus in The second half of the 20th century was the golden age of virus discovery and most of the over 2, recognised species of animal, plant, and bacterial viruses were discovered during these years.
In , the hepatitis B virus was discovered by Baruch Blumberg ,  and in , Howard Temin described the first retrovirus.
Viruses are found wherever there is life and have probably existed since living cells first evolved. This provides an invaluable source of information for paleovirologists to trace back ancient viruses that have existed up to millions of years ago.
There are three main hypotheses that aim to explain the origins of viruses: In the past, there were problems with all of these hypotheses: The escape hypothesis did not explain the complex capsids and other structures on virus particles.
The virus-first hypothesis contravened the definition of viruses in that they require host cells. The evidence for an ancestral world of RNA cells  and computer analysis of viral and host DNA sequences are giving a better understanding of the evolutionary relationships between different viruses and may help identify the ancestors of modern viruses.
To date, such analyses have not proved which of these hypotheses is correct. Opinions differ on whether viruses are a form of life, or organic structures that interact with living organisms.
Although they have genes, they do not have a cellular structure, which is often seen as the basic unit of life. Viruses do not have their own metabolism , and require a host cell to make new products.
They differ from autonomous growth of crystals as they inherit genetic mutations while being subject to natural selection. Virus self-assembly within host cells has implications for the study of the origin of life , as it lends further credence to the hypothesis that life could have started as self-assembling organic molecules.
Viruses display a wide diversity of shapes and sizes, called morphologies. In general, viruses are much smaller than bacteria.
Most viruses that have been studied have a diameter between 20 and nanometres. These are solutions of salts of heavy metals, such as tungsten , that scatter the electrons from regions covered with the stain.
When virions are coated with stain positive staining , fine detail is obscured. Negative staining overcomes this problem by staining the background only.
A complete virus particle, known as a virion, consists of nucleic acid surrounded by a protective coat of protein called a capsid. These are formed from identical protein subunits called capsomeres.
The capsid is made from proteins encoded by the viral genome and its shape serves as the basis for morphological distinction.
Complex viruses code for proteins that assist in the construction of their capsid. Proteins associated with nucleic acid are known as nucleoproteins , and the association of viral capsid proteins with viral nucleic acid is called a nucleocapsid.
The capsid and entire virus structure can be mechanically physically probed through atomic force microscopy. The poxviruses are large, complex viruses that have an unusual morphology.
The viral genome is associated with proteins within a central disc structure known as a nucleoid. The nucleoid is surrounded by a membrane and two lateral bodies of unknown function.
The virus has an outer envelope with a thick layer of protein studded over its surface. The whole virion is slightly pleiomorphic , ranging from ovoid to brick shape.
The capsid appears hexagonal under an electron microscope, therefore the capsid is probably icosahedral.
Provisionally named Megavirus chilensis , it can be seen with a basic optical microscope. Some viruses that infect Archaea have complex structures that are unrelated to any other form of virus, with a wide variety of unusual shapes, ranging from spindle-shaped structures, to viruses that resemble hooked rods, teardrops or even bottles.
Other archaeal viruses resemble the tailed bacteriophages, and can have multiple tail structures.
An enormous variety of genomic structures can be seen among viral species ; as a group, they contain more structural genomic diversity than plants, animals, archaea, or bacteria.
There are millions of different types of viruses,  although only about 5, types have been described in detail. The vast majority of viruses have RNA genomes.
Viral genomes are circular , as in the polyomaviruses , or linear , as in the adenoviruses. The type of nucleic acid is irrelevant to the shape of the genome.
Among RNA viruses and certain DNA viruses, the genome is often divided up into separate parts, in which case it is called segmented.
For RNA viruses, each segment often codes for only one protein and they are usually found together in one capsid.
All segments are not required to be in the same virion for the virus to be infectious, as demonstrated by brome mosaic virus and several other plant viruses.
A viral genome, irrespective of nucleic acid type, is almost always either single-stranded or double-stranded.
Single-stranded genomes consist of an unpaired nucleic acid, analogous to one-half of a ladder split down the middle. Double-stranded genomes consist of two complementary paired nucleic acids, analogous to a ladder.
The virus particles of some virus families, such as those belonging to the Hepadnaviridae , contain a genome that is partially double-stranded and partially single-stranded.
For most viruses with RNA genomes and some with single-stranded DNA genomes, the single strands are said to be either positive-sense called the plus-strand or negative-sense called the minus-strand , depending on if they are complementary to the viral messenger RNA mRNA.
Positive-sense viral RNA is in the same sense as viral mRNA and thus at least a part of it can be immediately translated by the host cell. Genome size varies greatly between species.
In general, RNA viruses have smaller genome sizes than DNA viruses because of a higher error-rate when replicating, and have a maximum upper size limit.
In contrast, DNA viruses generally have larger genomes because of the high fidelity of their replication enzymes.
Viruses undergo genetic change by several mechanisms. This can be a result of recombination or reassortment. When this happens with influenza viruses, pandemics might result.
Such quasispecies are a prime target for natural selection. Segmented genomes confer evolutionary advantages; different strains of a virus with a segmented genome can shuffle and combine genes and produce progeny viruses or offspring that have unique characteristics.
This is called reassortment or viral sex. Genetic recombination is the process by which a strand of DNA is broken and then joined to the end of a different DNA molecule.
This can occur when viruses infect cells simultaneously and studies of viral evolution have shown that recombination has been rampant in the species studied.
Viral populations do not grow through cell division, because they are acellular. Instead, they use the machinery and metabolism of a host cell to produce multiple copies of themselves, and they assemble in the cell.
The life cycle of viruses differs greatly between species but there are six basic stages in the life cycle of viruses: Attachment is a specific binding between viral capsid proteins and specific receptors on the host cellular surface.
This specificity determines the host range of a virus. For example, HIV infects a limited range of human leucocytes. This mechanism has evolved to favour those viruses that infect only cells in which they are capable of replication.
Attachment to the receptor can induce the viral envelope protein to undergo changes that results in the fusion of viral and cellular membranes, or changes of non-enveloped virus surface proteins that allow the virus to enter.
Virions enter the host cell through receptor-mediated endocytosis or membrane fusion. This is often called viral entry.
The infection of plant and fungal cells is different from that of animal cells. Plants have a rigid cell wall made of cellulose , and fungi one of chitin, so most viruses can get inside these cells only after trauma to the cell wall.
Given that bacterial cell walls are much thinner than plant cell walls due to their much smaller size, some viruses have evolved mechanisms that inject their genome into the bacterial cell across the cell wall, while the viral capsid remains outside.
Uncoating is a process in which the viral capsid is removed: This may be by degradation by viral enzymes or host enzymes or by simple dissociation; the end-result is the releasing of the viral genomic nucleic acid.
Replication of viruses involves primarily multiplication of the genome. Replication involves synthesis of viral messenger RNA mRNA from "early" genes with exceptions for positive sense RNA viruses , viral protein synthesis , possible assembly of viral proteins, then viral genome replication mediated by early or regulatory protein expression.
This may be followed, for complex viruses with larger genomes, by one or more further rounds of mRNA synthesis: Assembly — Following the structure-mediated self- assembly of the virus particles, some modification of the proteins often occurs.
In viruses such as HIV, this modification sometimes called maturation occurs after the virus has been released from the host cell. Release — Viruses can be released from the host cell by lysis , a process that kills the cell by bursting its membrane and cell wall if present: This is a feature of many bacterial and some animal viruses.
Some viruses undergo a lysogenic cycle where the viral genome is incorporated by genetic recombination into a specific place in the host's chromosome.
The viral genome is then known as a " provirus " or, in the case of bacteriophages a " prophage ". The viral genome is mostly silent within the host.
At some point, the provirus or prophage may give rise to active virus, which may lyse the host cells. During this process the virus acquires its envelope, which is a modified piece of the host's plasma or other, internal membrane.
The genetic material within virus particles, and the method by which the material is replicated, varies considerably between different types of viruses.
The range of structural and biochemical effects that viruses have on the host cell is extensive.
The causes of death include cell lysis, alterations to the cell's surface membrane and apoptosis. Some viruses, such as Epstein—Barr virus , can cause cells to proliferate without causing malignancy,  while others, such as papillomaviruses , are established causes of cancer.
Some viruses cause no apparent changes to the infected cell. Cells in which the virus is latent and inactive show few signs of infection and often function normally.
This is often the case with herpes viruses. Viruses are by far the most abundant biological entities on Earth and they outnumber all the others put together.
Other viruses, such as rabies virus, can infect different species of mammals and are said to have a broad range.
Classification seeks to describe the diversity of viruses by naming and grouping them on the basis of similarities. Viruses were grouped according to their shared properties not those of their hosts and the type of nucleic acid forming their genomes.
The system proposed by Lwoff, Horne and Tournier was never fully accepted by the ICTV because small genome size viruses and their high rate of mutation makes it difficult to determine their ancestry beyond order.
As such, the Baltimore classification is used to supplement the more traditional hierarchy. The International Committee on Taxonomy of Viruses ICTV developed the current classification system and wrote guidelines that put a greater weight on certain virus properties to maintain family uniformity.
A unified taxonomy a universal system for classifying viruses has been established. Only a small part of the total diversity of viruses has been studied.
As of , nine orders, families, 46 subfamilies , genera, and 4, species of viruses have been defined by the ICTV.
The Baltimore classification of viruses is based on the mechanism of mRNA production. Viruses must generate mRNAs from their genomes to produce proteins and replicate themselves, but different mechanisms are used to achieve this in each virus family.
This classification places viruses into seven groups:. As an example of viral classification, the chicken pox virus, varicella zoster VZV , belongs to the order Herpesvirales , family Herpesviridae , subfamily Alphaherpesvirinae , and genus Varicellovirus.
The complete set of viruses in an organism or habitat is called the virome ; for example, all human viruses constitute the human virome. Examples of common human diseases caused by viruses include the common cold , influenza, chickenpox , and cold sores.
The relative ability of viruses to cause disease is described in terms of virulence. Other diseases are under investigation to discover if they have a virus as the causative agent, such as the possible connection between human herpesvirus 6 HHV6 and neurological diseases such as multiple sclerosis and chronic fatigue syndrome.
Viruses have different mechanisms by which they produce disease in an organism, which depends largely on the viral species.
Mechanisms at the cellular level primarily include cell lysis, the breaking open and subsequent death of the cell. In multicellular organisms , if enough cells die, the whole organism will start to suffer the effects.
Although viruses cause disruption of healthy homeostasis , resulting in disease, they may exist relatively harmlessly within an organism. An example would include the ability of the herpes simplex virus , which causes cold sores, to remain in a dormant state within the human body.
This is called latency  and is a characteristic of the herpes viruses, including Epstein—Barr virus, which causes glandular fever, and varicella zoster virus , which causes chickenpox and shingles.
Most people have been infected with at least one of these types of herpes virus. Some viruses can cause lifelong or chronic infections, where the viruses continue to replicate in the body despite the host's defence mechanisms.
People chronically infected are known as carriers, as they serve as reservoirs of infectious virus. Viral epidemiology is the branch of medical science that deals with the transmission and control of virus infections in humans.
Transmission of viruses can be vertical, which means from mother to child, or horizontal, which means from person to person. Examples of vertical transmission include hepatitis B virus and HIV, where the baby is born already infected with the virus.
Horizontal transmission is the most common mechanism of spread of viruses in populations. Transmission can occur when: The rate or speed of transmission of viral infections depends on factors that include population density, the number of susceptible individuals, i.
Epidemiology is used to break the chain of infection in populations during outbreaks of viral diseases.
It is important to find the source, or sources, of the outbreak and to identify the virus. Once the virus has been identified, the chain of transmission can sometimes be broken by vaccines.
When vaccines are not available, sanitation and disinfection can be effective. Often, infected people are isolated from the rest of the community, and those that have been exposed to the virus are placed in quarantine.
If outbreaks spread worldwide, they are called pandemics. Native American populations were devastated by contagious diseases, in particular, smallpox , brought to the Americas by European colonists.
The damage done by this disease significantly aided European attempts to displace and conquer the native population.
A pandemic is a worldwide epidemic. The flu pandemic , which lasted until , was a category 5 influenza pandemic caused by an unusually severe and deadly influenza A virus.
The victims were often healthy young adults, in contrast to most influenza outbreaks, which predominantly affect juvenile, elderly, or otherwise-weakened patients.
Most researchers believe that HIV originated in sub-Saharan Africa during the 20th century;  it is now a pandemic, with an estimated Several highly lethal viral pathogens are members of the Filoviridae.
Filoviruses are filament-like viruses that cause viral hemorrhagic fever , and include ebolaviruses and marburgviruses. Marburg virus , first discovered in , attracted widespread press attention in April for an outbreak in Angola.
The worst and most recent one is the West Africa epidemic. Viruses are an established cause of cancer in humans and other species.
Viral cancers occur only in a minority of infected persons or animals. Cancer viruses come from a range of virus families, including both RNA and DNA viruses, and so there is no single type of " oncovirus " an obsolete term originally used for acutely transforming retroviruses.
The development of cancer is determined by a variety of factors such as host immunity  and mutations in the host.
The most recently discovered human cancer virus is a polyomavirus Merkel cell polyomavirus that causes most cases of a rare form of skin cancer called Merkel cell carcinoma.
The body's first line of defence against viruses is the innate immune system. This comprises cells and other mechanisms that defend the host from infection in a non-specific manner.
This means that the cells of the innate system recognise, and respond to, pathogens in a generic way, but, unlike the adaptive immune system , it does not confer long-lasting or protective immunity to the host.
RNA interference is an important innate defence against viruses. When such a virus infects a cell, it releases its RNA molecule or molecules, which immediately bind to a protein complex called a dicer that cuts the RNA into smaller pieces.
Rotaviruses have evolved to avoid this defence mechanism by not uncoating fully inside the cell, and releasing newly produced mRNA through pores in the particle's inner capsid.
Their genomic dsRNA remains protected inside the core of the virion. When the adaptive immune system of a vertebrate encounters a virus, it produces specific antibodies that bind to the virus and often render it non-infectious.
This is called humoral immunity. Two types of antibodies are important. The first, called IgM , is highly effective at neutralising viruses but is produced by the cells of the immune system only for a few weeks.
The second, called IgG , is produced indefinitely. The presence of IgM in the blood of the host is used to test for acute infection, whereas IgG indicates an infection sometime in the past.
Antibodies can continue to be an effective defence mechanism even after viruses have managed to gain entry to the host cell.
A protein that is in cells, called TRIM21 , can attach to the antibodies on the surface of the virus particle. This primes the subsequent destruction of the virus by the enzymes of the cell's proteosome system.
A second defence of vertebrates against viruses is called cell-mediated immunity and involves immune cells known as T cells. The body's cells constantly display short fragments of their proteins on the cell's surface, and, if a T cell recognises a suspicious viral fragment there, the host cell is destroyed by killer T cells and the virus-specific T-cells proliferate.
Cells such as the macrophage are specialists at this antigen presentation. This is a hormone produced by the body when viruses are present. Its role in immunity is complex; it eventually stops the viruses from reproducing by killing the infected cell and its close neighbours.
Not all virus infections produce a protective immune response in this way. HIV evades the immune system by constantly changing the amino acid sequence of the proteins on the surface of the virion.
This is known as "escape mutation" as the viral epitopes escape recognition by the host immune response. These persistent viruses evade immune control by sequestration, blockade of antigen presentation , cytokine resistance, evasion of natural killer cell activities, escape from apoptosis , and antigenic shift.
Because viruses use vital metabolic pathways within host cells to replicate, they are difficult to eliminate without using drugs that cause toxic effects to host cells in general.
The most effective medical approaches to viral diseases are vaccinations to provide immunity to infection, and antiviral drugs that selectively interfere with viral replication.
Vaccination is a cheap and effective way of preventing infections by viruses. Vaccines were used to prevent viral infections long before the discovery of the actual viruses.
Their use has resulted in a dramatic decline in morbidity illness and mortality death associated with viral infections such as polio , measles , mumps and rubella.
Such viruses are called attenuated. Live vaccines can be dangerous when given to people with a weak immunity who are described as immunocompromised , because in these people, the weakened virus can cause the original disease.
These vaccines use only the capsid proteins of the virus. Hepatitis B vaccine is an example of this type of vaccine. Antiviral drugs are often nucleoside analogues fake DNA building-blocks , which viruses mistakenly incorporate into their genomes during replication.
The life-cycle of the virus is then halted because the newly synthesised DNA is inactive. This is because these analogues lack the hydroxyl groups, which, along with phosphorus atoms, link together to form the strong "backbone" of the DNA molecule.
This is called DNA chain termination. Aciclovir is one of the oldest and most frequently prescribed antiviral drugs.
HIV is dependent on a proteolytic enzyme called the HIV-1 protease for it to become fully infectious. There is a large class of drugs called protease inhibitors that inactivate this enzyme.
Hepatitis C is caused by an RNA virus. There is now an effective treatment that uses the nucleoside analogue drug ribavirin combined with interferon.
Viruses infect all cellular life and, although viruses occur universally, each cellular species has its own specific range that often infect only that species.
Viruses are important pathogens of livestock. Diseases such as foot-and-mouth disease and bluetongue are caused by viruses. Canine parvovirus is caused by a small DNA virus and infections are often fatal in pups.
There are many types of plant virus, but often they cause only a loss of yield , and it is not economically viable to try to control them.
Plant viruses are often spread from plant to plant by organisms , known as vectors. These are normally insects, but some fungi, nematode worms , and single-celled organisms have been shown to be vectors.
When control of plant virus infections is considered economical, for perennial fruits, for example, efforts are concentrated on killing the vectors and removing alternate hosts such as weeds.
Plants have elaborate and effective defence mechanisms against viruses. One of the most effective is the presence of so-called resistance R genes.
Each R gene confers resistance to a particular virus by triggering localised areas of cell death around the infected cell, which can often be seen with the unaided eye as large spots.
This stops the infection from spreading. Plant virus particles or virus-like particles VLPs have applications in both biotechnology and nanotechnology.
The capsids of most plant viruses are simple and robust structures and can be produced in large quantities either by the infection of plants or by expression in a variety of heterologous systems.
Plant virus particles can be modified genetically and chemically to encapsulate foreign material and can be incorporated into supramolecular structures for use in biotechnology.
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, in the case of the T4 phage , in just over twenty minutes after injection over three hundred phages could be released.
The major way 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.
Some viruses replicate within archaea: 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.
The organic molecules released from the dead bacterial cells stimulate fresh bacterial and algal growth, in a process known as the viral shunt.
In January , scientists reported that million viruses, mainly of marine origin, are deposited daily from the Earth 's atmosphere onto every square meter of the planet's surface, as the result of a global atmospheric stream of viruses, circulating above the weather system, but below the altitude of usual airline travel, distributing viruses around the planet.
Like any organism, marine mammals are susceptible to viral infections. In and , thousands of harbour seals were killed in Europe by phocine distemper virus.
Viruses are an important natural means of transferring genes between different species, which increases genetic diversity and drives evolution.
Viruses are important to the study of molecular and cell biology as they provide simple systems that can be used to manipulate and investigate the functions of cells.
Geneticists often use viruses as vectors to introduce genes into cells that they are studying. This is useful for making the cell produce a foreign substance, or to study the effect of introducing a new gene into the genome.
In similar fashion, virotherapy uses viruses as vectors to treat various diseases, as they can specifically target cells and DNA.
It shows promising use in the treatment of cancer and in gene therapy. Eastern European scientists have used phage therapy as an alternative to antibiotics for some time, and interest in this approach is increasing, because of the high level of antibiotic resistance now found in some pathogenic bacteria.
Industrial processes have been recently developed using viral vectors and a number of pharmaceutical proteins are currently in pre-clinical and clinical trials.
Virotherapy involves the use of genetically modified viruses to treat diseases. Talimogene laherparepvec T-VEC , for example, is a modified herpes simplex virus that has had a gene, which is required for viruses to replicate in healthy cells, deleted and replaced with a human gene GM-CSF that stimulates immunity.
When this virus infects cancer cells, it destroys them and in doing so the presence the GM-CSF gene attracts dendritic cells from the surrounding tissues of the body.
The dendritic cells process the dead cancer cells and present components of them to other cells of the immune system. Current trends in nanotechnology promise to make much more versatile use of viruses.
From the viewpoint of a materials scientist, viruses can be regarded as organic nanoparticles. Their surface carries specific tools designed to cross the barriers of their host cells.
The size and shape of viruses, and the number and nature of the functional groups on their surface, is precisely defined.
As such, viruses are commonly used in materials science as scaffolds for covalently linked surface modifications. A particular quality of viruses is that they can be tailored by directed evolution.
The powerful techniques developed by life sciences are becoming the basis of engineering approaches towards nanomaterials, opening a wide range of applications far beyond biology and medicine.
Because of their size, shape, and well-defined chemical structures, viruses have been used as templates for organising materials on the nanoscale.
In this application, the virus particles separate the fluorescent dyes used for signalling to prevent the formation of non-fluorescent dimers that act as quenchers.
After a virus devastates the global human population, survivors in Antarctica desperately try to find a cure and save the human race.
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