Portal:Viruses/Selected article

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Dengue fever, or breakbone fever, is an infectious tropical disease caused by the dengue virus. Symptoms include fever, headache, muscle and joint pains, and a characteristic skin rash that is similar to measles. Rarely, the disease develops into the life-threatening dengue haemorrhagic fever, resulting in bleeding, low levels of blood platelets and blood plasma leakage, or into dengue shock syndrome, where dangerously low blood pressure occurs.

Dengue is transmitted by several species of Aedes mosquito, mainly A. aegypti. First described in 1779, the incidence of dengue fever has increased dramatically since the Second World War, with around 390 million people infected each year, of whom about 40,000 die. The virus is endemic in more than 120 countries, mainly in Southeast Asia, South Asia and South America. Infection with one of the five viral types usually gives lifelong immunity to that type but only short-term immunity to the others, and subsequent infection with a different type increases the risk of severe complications. Treatment of acute dengue is supportive, using oral or intravenous fluids for mild or moderate disease; blood transfusion may be required in more severe cases. A vaccine against dengue fever is available. Reducing mosquito habitats and limiting exposure to bites are the other main methods of control.

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Polio, also called poliomyelitis or infantile paralysis, was one of the most feared childhood diseases of the 20th century. Poliovirus, the causative agent, only naturally infects humans and spreads via the faecal–oral route. Most infections cause no or minor symptoms. In around 1% of cases, the virus enters the central nervous system, causing aseptic meningitis. There it can preferentially infect and destroy motor neurons, leading in 0.1–0.5% of cases to muscle weakness and acute flaccid paralysis. Spinal polio accounts for nearly 80% of paralytic cases, with asymmetric paralysis of the legs being typical; in a quarter of these cases permanent severe disability results. Bulbar involvement is rare, but in severe cases the virus can prevent breathing by affecting the phrenic nerve, so that patients require mechanical ventilation with an iron lung or similar device.

Depictions in ancient art show that the disease has existed for thousands of years. The virus was an endemic pathogen until the 1880s, when major epidemics began to occur in Europe and later the United States. Polio vaccines were developed in the 1950s and a global eradication campaign started in 1988. The annual incidence of wild-type disease fell from many hundreds of thousands to 22 in 2017, but has since resurged to a few hundreds.

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Smallpox is an eradicated infectious disease of humans caused by the Variola major and V. minor viruses. V. major causes a serious disease with a mortality rate of around 30%; V. minor is associated with much milder symptoms and mortality below 1%. The virus is mainly transmitted by the respiratory route but can also be carried on contaminated objects. Smallpox preferentially infects skin cells, resulting in a usually maculopapular rash, and later, raised fluid-filled blisters. Most V. major survivors have permanent scarring, commonly on the face, which can be extensive. Less common long-term complications include blindness resulting from corneal ulceration and scarring, and in young children, limb deformities due to arthritis and osteomyelitis.

Smallpox probably emerged in human populations in about 10,000 BC; the mummified body of Egyptian pharaoh Ramses V shows evidence of smallpox rash. The disease was responsible for an estimated 300–500 million deaths during the 20th century. Smallpox vaccine, the earliest vaccine, was developed in the 18th century. After intensive vaccination campaigns, the last natural infection occurred in 1977. Smallpox was certified the first infectious disease to be eradicated globally in 1979. Debate is ongoing over whether all stocks of the virus should be destroyed.

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Influenza, or flu, is an infectious disease caused by some orthomyxoviruses, that affects birds and some mammals including humans, horses and pigs. Influenza is predominantly transmitted through the air by coughs or sneezes, creating aerosols containing the virus. It can also be transmitted by contact with bird droppings or nasal secretions, or by touching contaminated surfaces. As the virus can be inactivated by soap, frequent hand washing reduces the risk of infection. Around a third of cases show no symptoms. The most common symptoms include fever, runny nose, sore throat, muscle pains, headache, cough and fatigue. Influenza is occasionally associated with nausea and vomiting, particularly in children. Pneumonia is a rare complication which can be life-threatening.

Influenza spreads around the world in seasonal epidemics, resulting in about 3–5 million cases of severe illness annually, and about 250,000–500,000 deaths, mainly in the young, the old and those with other health problems. Annual influenza vaccinations are recommended for those at high risk. Sporadic influenza pandemics have been recorded since at least the 16th century. The Spanish flu pandemic of 1918–20 is estimated to have killed 50–100 million people.

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Hepatitis B is an infectious inflammatory disease of the liver caused by the hepatitis B virus (HBV), a hepadnavirus. It affects humans and possibly other great apes. The virus is transmitted by exposure to infectious blood or some body fluids. Mother-to-child transmission is a major route in endemic countries. HBV is 50–100 times more infectious than HIV. The virus replicates in liver cells, and enters the blood where viral proteins and antiviral antibodies are found.

Acute infection is often asymptomatic but can cause liver inflammation resulting in vomiting, jaundice and, rarely, death. Over 95% of infected adults and older children clear the infection spontaneously, developing protective immunity. Only 30% of children aged 1–6 years and 5% of newborns infected perinatally clear the infection. Chronic hepatitis B may eventually progress to cirrhosis and liver cancer, causing death in around 40% of those chronically infected. The virus has infected humans since at least the Bronze Age, with HBV DNA being found in 4,500-year-old human remains. About a third of the global population has been infected at one point in their lives, including nearly 350 million who are chronic carriers. The virus is endemic in East Asia and sub-Saharan Africa. Infection can be prevented by vaccination.

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Herpes simplex is caused by herpes simplex virus types 1 and 2 of the Herpesviridae family, with 60–95% of adults being infected with one of the types. Common forms of infection are oral herpes, which can result in cold sores, and genital herpes. Active disease often involves blisters containing infectious virus, although the genital form is frequently asymptomatic. Less common disorders associated with the viruses include herpetic whitlow, herpes gladiatorum, ocular herpes, herpesviral encephalitis and Mollaret's meningitis.

After initial infection, virus particles are transported along sensory nerves to the cell bodies in the ganglion, where they become latent and remain lifelong. Periods of remission alternate with outbreaks of active disease, in which the virus multiplies in the nerve cell and new virus particles are transported along the nerve fibre to the nerve terminals in the skin, where they are released. What causes these recurrences is unclear. Transmission is usually by direct contact with a lesion or with body fluids, and can occur during periods of asymptomatic shedding. Neonatal herpes is possible after transmission from the mother. Barrier protection methods reduce genital herpes risk. No vaccine or cure exists, but antiviral treatment can alleviate symptoms and reduce viral shedding.

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The common cold is an upper respiratory tract disease that mainly affects the nose, and sometimes the throat, larynx and sinuses. Over 200 viruses can cause colds, most commonly rhinoviruses but also coronaviruses, influenza viruses, adenoviruses and others. Adults catch an average of 2–3 colds a year and children 6–8, making it the most common infectious human disease. The economic costs are huge, with colds responsible for 40% of time lost from work in the U.S. Colds are described in the Egyptian Ebers papyrus, the oldest surviving medical text, written before the 16th century BCE.

Symptoms include cough, sore throat, runny nose, nasal congestion, sneezing and sometimes muscle aches and headache; fever is common in young children. Symptoms typically resolve in 7–10 days, although some can last up to 3 weeks. The immune response to infection, rather than tissue destruction by the virus, causes most of the symptoms. Transmission occurs via airborne droplets and by contact with nasal secretions or contaminated objects. The viruses that cause colds can survive for prolonged periods in the environment (over 18 hours for rhinoviruses). Hand washing can help to prevent spread. No effective antiviral treatment or vaccine currently exists.

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Shingles, or herpes zoster, is a painful skin rash with blisters that, characteristically, occurs in a stripe limited to just one side of the body. The rash usually heals within 2–5 weeks, but around one in five people experience residual nerve pain for months or years.

Shingles is caused by varicella zoster virus (VZV), an alpha-herpesvirus. Initial VZV infection usually occurs in childhood causing chickenpox. After this resolves, the virus is not eliminated from the body, but remains latent in the nerve cell bodies of the dorsal root or trigeminal ganglia, without causing symptoms. Years or decades later, shingles occurs when virions in a single ganglion reactivate, travel down nerve fibres and infect the skin around the nerve. The shingles rash is restricted to the area of skin supplied by a single spinal nerve, termed the dermatome. Exactly how VZV remains latent in the body, and subsequently reactivates, is unclear.

Around a third of the population will develop shingles. Repeated episodes are rare. In the United States, about half the cases occur in people aged 50 years or older. Vaccination at least halves the risk, and prompt treatment with aciclovir or related antiviral drugs can reduce the severity and duration of the rash.

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Diagram showing the three layers of the meninges: the dura mater (blue), arachnoid mater (green) and pia mater (fawn)

Meningitis is an acute inflammation of the meninges, protective membranes covering the brain and spinal cord. Symptoms in adults include headache, fever and neck stiffness, as well as sometimes confusion or altered consciousness, vomiting, and an inability to tolerate light or loud noises. Children often show only nonspecific symptoms, such as irritability or drowsiness.

The most common cause is infection with viruses including enteroviruses, herpes simplex virus (mainly HSV-2), varicella zoster virus, mumps virus, HIV and lymphocytic choriomeningitis. In Western countries, viral meningitis occurs in around 11 people per 100,000 each year. Infection with bacteria, fungi, protozoa and parasites can also cause meningitis, and there are several non-infectious causes. Although some forms of meningitis can be life-threatening, viral meningitis is generally more benign than that caused by bacterial infection. It usually resolves spontaneously and is rarely fatal. HSV-2 can cause a chronic, recurrent form called Mollaret's meningitis.

Polymerase chain reaction of cerebrospinal fluid and identification of antibodies can be used to differentiate between viral causes. Viral meningitis typically only requires supportive therapy; meningitis caused by HSV or varicella zoster virus sometimes responds to treatment with antiviral drugs such as aciclovir. Mumps-associated meningitis can be prevented by vaccination.

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Hepatitis C is an infectious disease affecting mainly the liver, caused by hepatitis C virus (HCV), an RNA virus of the Flaviviridae family which only infects humans and chimpanzees. A "non-A non-B hepatitis" was postulated in the 1970s, and HCV was demonstrated in 1989. HCV is spread primarily by blood-to-blood contact associated with intravenous drug use in the developed world, and with improperly sterilised medical equipment and blood transfusions in the developing world. In about 80% of those infected, the virus establishes a chronic infection in the liver, and around 10–30% of those infected will develop cirrhosis over 30 years. Some people with cirrhosis go on to develop liver failure, liver cancer or other serious complications.

An estimated 143 million people worldwide (2%) have chronic HCV infections as of 2015. The prevalence is highest in Central and East Asia, North Africa and the Middle East. The virus causes around a quarter of cases of cirrhosis and hepatocellular carcinoma, and is a major reason for liver transplantation. The recommended therapy is an appropriate combination of protease inhibitors. Although 95% of people treated in this way are cured, the treatments are expensive and older therapies are less effective. No vaccine against hepatitis C is available.

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Gastroenteritis is an inflammation of the gastrointestinal tract involving both the stomach and small intestine, which results in diarrhoea and vomiting, and sometimes abdominal pain. It is usually caused by a virus: most commonly rotavirus and norovirus, but also adenovirus and astrovirus. Other major infectious causes include Campylobacter, Escherichia coli, Vibrio cholerae and some other bacteria, as well as protozoa. Viruses, particularly rotavirus, cause about 70% of gastroenteritis episodes in children, while norovirus is the leading cause of gastroenteritis among adults in America, causing over 90% of outbreaks.

Transmission can be from consumption of improperly prepared foods or contaminated water, or by close contact with infectious individuals. Good sanitation practices and a convenient supply of uncontaminated water are important for reducing infection. Personal measures such as hand washing with soap can decrease incidence by as much as 30%. An estimated 2 billion cases of gastroenteritis occurred globally in 2015, mainly among children and people in developing countries, resulting in 1.3 million deaths. Gastroenteritis is usually an acute and self-limiting disease that does not require medication; the main treatment is rehydration using oral rehydration therapy. A rotavirus vaccine is available.

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Yellow fever is an acute haemorrhagic fever caused by the yellow fever virus, an RNA virus in the Flaviviridae family. It infects humans, other primates, and Aedes aegypti and other mosquito species, which act as the vector. After transmission by the bite of a female mosquito, the virus replicates in lymph nodes, infecting dendritic cells, and can then spread to liver hepatocytes. Symptoms generally last 3–4 days, and include fever, nausea and muscle pain. In around 15% of people, a toxic phase follows with recurring fever, liver damage and jaundice, sometimes accompanied by bleeding and kidney failure; death occurs in 20–50% of those who develop jaundice. Infection otherwise leads to lifelong immunity.

The first definitive outbreak of yellow fever was in Barbados in 1647, and major epidemics have occurred in the Americas and southern Europe since that date. Yellow fever is endemic in tropical and subtropical areas of South America and Africa; its incidence has been increasing since the 1980s. An estimated 200,000 cases and 30,000 deaths occur each year, with almost 90% of cases being in Africa. Antiviral therapy is not effective. A vaccine is available, and vaccination, mosquito control and bite prevention are the main preventive measures.

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Chickenpox is caused by initial infection with varicella zoster virus, a DNA virus of the Alphaherpesvirinae subfamily. The virus naturally infects only humans, but some other primates have been infected artificially. Symptoms appear 10–21 days after exposure: an itchy vesicular skin rash, and small ulcers in the oral cavity and tonsil areas. The rash usually resolves by 7 days, but the virus remains latent in nerve cell bodies, and can emerge years or decades later to cause shingles. Chickenpox is transmitted by the respiratory route, as well as direct contact with lesions.

A classic disease of childhood, the highest prevalence occurs at 4–10 years. Chickenpox is rarely fatal in people with a normal immune system, with around 6,400 deaths worldwide in 2015, about 1 in 60,000 infections. Adults often have more severe symptoms than children, and are at higher risk of complications such as pneumonia, bronchitis, hepatitis and encephalitis. Pregnant women and people with a suppressed immune system have the highest complication risk. Chickenpox during the first 28 weeks of gestation can lead to foetal malformations. Infection in adults is usually treated with antiviral drugs, such as aciclovir or valaciclovir, which reduces symptom severity and the risk of complications. A vaccine is available.

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AIDS is a progressive immunodeficiency disease first recognised in 1981, caused by infection with human immunodeficiency virus (HIV). Initial infection is sometimes associated with a brief influenza-like illness, followed by a period of 3–20 years without symptoms during which the immune system deteriorates. The individual becomes highly susceptible to common infections such as tuberculosis, as well as opportunistic infections, such as Pneumocystis pneumonia, and tumours, such as Kaposi's sarcoma and lymphoma. Severe weight loss is also a feature of AIDS. Without treatment, the average survival time after infection is around 9–11 years, depending on the HIV subtype.

HIV/AIDS is a global pandemic. As of 2018, nearly 38 million people worldwide are estimated to be infected with HIV, with 1.8 million new infections occurring each year. AIDS has caused around 32 million deaths. The major routes of transmission are unsafe heterosexual or male/male sex, contaminated needles and blood transfusions, and from mother to child, either at birth or via breastfeeding. Combination antiretroviral therapy does not eliminate the virus, but delays progression to AIDS and can lead to a near-normal life expectancy. Drugs to prevent infection are now available, but there is no effective HIV vaccine.

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Cervical cancer is a tumour of the cervix, the part of the uterus that adjoins the vagina in the female reproductive tract. Certain types of human papillomavirus (HPV) are implicated in more than 90% of these cancers, although the great majority of HPV infections of the cervix are not associated with cancer. HPV is transmitted by vaginal sex, infecting cervical epithelial cells. In a minority of cases, infection persists for years, and pre-cancerous changes called cervical intraepithelial neoplasia can develop, which sometimes progress to cancer. Squamous cell carcinomas are the most common. HPV infection is considered essential for nearly all forms of cervical cancer to develop, but other risk factors are also involved, including smoking, HIV infection and other forms of immune suppression, and long-term use of oral contraceptives.

Cervical cancer is the fourth most common cancer in women worldwide. It can be detected by screening; screening every 3–5 years, with appropriate follow-up, can reduce cancer incidence by up to 80%. HPV vaccines protect against high-risk virus strains, and can prevent up to 90% of cervical cancers. Where screening and vaccination are not available, cervical cancer has substantial mortality; worldwide, an estimated 569,000 cases and 311,000 deaths occurred in 2018, with around 85% of cases being in developing countries.

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Ebola virus disease is a viral haemorrhagic fever of humans and other primates caused by ebolaviruses, RNA viruses in the Filoviridae family. The natural host is thought to be megabats. The virus spreads by contact with body fluids, such as blood. After an incubation period of 2–21 days, nonspecific symptoms including fever, sore throat, muscle pain and headache develop, followed by vomiting, diarrhoea, rash, liver and kidney impairment, and sometimes internal or external bleeding. On average, around half of those infected die, often from hypovolaemic shock. The virus sometimes persists in semen or breast milk for several weeks or months after recovery.

The disease was first identified in 1976 by Peter Piot and others in two simultaneous outbreaks, one in Nzara, Sudan and the other in Yambuku, then in Zaire, near the Ebola River, for which the disease is named. Outbreaks have occurred intermittently in tropical regions of sub-Saharan Africa, with a few thousand cases documented before the epidemic in Western Africa of 2013–16, in which nearly 29,000 cases with more than 11,000 deaths were reported. More than 3000 cases were documented in a subsequent epidemic in the Democratic Republic of the Congo in 2018–20, with more than 2000 deaths. Prevention of outbreaks focuses on limiting virus transmission from infected animals to humans, for example, by taking precautions when handling bush meat. Ebola vaccines are now available. Monoclonal antibody treatments can improve outcomes, and a combination of three monoclonals has been licensed.

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Variant Creutzfeldt–Jakob disease, or vCJD, is a rare type of central nervous system disease within the transmissible spongiform encephalopathy family, caused by a prion. First identified in 1996, vCJD is now distinguished from classic CJD. The incubation period is believed to be years, possibly over 50 years. Prion protein can be detected in appendix and lymphoid tissue (pictured) up to two years before the onset of neurological symptoms, which include psychiatric problems, behavioural changes and painful sensations. Abnormal prion proteins build up as amyloid deposits in the brain, which acquires a characteristic spongiform appearance, with many round vacuoles in the cerebellum and cerebrum. The average life expectancy after symptoms start is 13 months.

About 170 cases have been recorded in the UK, and 50 cases in the rest of the world. The estimated prevalence in the UK is about 1 in 2000, higher than the reported cases. Transmission is believed to be mainly from consuming beef contaminated with the bovine spongiform encephalopathy prion, but may potentially also occur via blood products or contaminated surgical equipment. Infection is also believed to require a specific genetic susceptibility in the PRNP-encoding gene. Human PRNP protein can have either methionine or valine at position 129; nearly all of those affected had two copies of the methionine-containing form, found in 40% of Caucasians.

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Measles is a disease that only affects humans caused by the measles virus, an RNA virus in the Paramyxoviridae family. It is highly contagious, with transmission occurring via the respiratory route or by contact with secretions. Symptoms generally develop 10–12 days after exposure and last 7–10 days; they include high fever, cough, rhinitis and conjunctivitis, white Koplik's spots inside the mouth and a generalised red maculopapular rash. Complications including diarrhoea, otitis media and pneumonia are relatively common; more rarely seizures, encephalitis, croup, corneal ulceration and blindness can occur. The risk of death is usually around 0.2%, but may be as high as 10–28% in areas with high levels of malnutrition and poor healthcare.

Measles was first described by Rhazes (860–932). The disease is estimated to have killed around 200 million people between 1855 and 2005. It affects about 20 million people a year, primarily in the developing areas of Africa and Asia, and is one of the leading vaccine-preventable disease causes of death. No antiviral drug is licensed. An effective measles vaccine is available, but uptake has been reduced by anti-vaccination campaigns, particularly the fraudulent claim that the MMR vaccine might be associated with autism. Rates of disease and deaths increased from 2017 to 2019, attributed to a decrease in immunisation.

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Foot-and-mouth disease or FMD is an economically important disease of even-toed ungulates (cloven-hoofed animals) and some other mammals caused by the FMD virus, a picornavirus. Hosts include cattle, water buffalo, sheep, goats, pigs, antelope, deer and bison; human infection is extremely rare. After a 1–12-day incubation, animals develop high fever, and then blisters inside the mouth (pictured) and on the hooves, which can rupture and cause lameness. Weight loss and reduction in milk production are other possible long-term consequences. Mortality in adult animals is low (2–5%). The virus is highly infectious, with transmission occurring via direct contact, aerosols, semen, consumption of infected food scraps or feed supplements, and via inanimate objects including fodder, farming equipment, vehicles, standing water, and the clothes and skin of humans. Some infected ruminants can transmit infection as asymptomatic carriers.

Friedrich Loeffler showed the disease to be viral in 1897. FMD was widely distributed in 1945. By 2014, North America, Australia, New Zealand, much of Europe, and some South American countries were free of the disease. Major outbreaks include one in the UK in 2001 that cost an estimated £8 billion. The virus is highly variable, with seven serotypes. A vaccine is available, but protection is temporary and strain specific. Other control methods include monitoring programmes, trade restrictions, quarantine, and the slaughter of infected and healthy at-risk animals.

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Rabies is a disease of humans and some other mammals, generally caused by the rabies virus, an RNA virus in the Rhabdoviridae family. A few cases have involved the closely related Australian bat lyssavirus. Rabies virus has a wide host range. Globally, dogs are the main source of human infections, with bats being important in the Americas; other naturally infected animals include monkeys, raccoons, foxes, skunks, cattle, horses, wolves, coyotes, cats, mongooses, bears, groundhogs, weasels and other carnivores. Transmission is commonly via saliva, usually but not always from bites; it can potentially occur via aerosols contacting mucous membranes. The typical human incubation period is 1–3 months. The neurotropic virus travels along neural pathways into the CNS and brain, where it causes meningoencephalitis. Nonspecific symptoms such as fever and headache are followed by neurological symptoms, including partial paralysis, confusion, agitation, paranoia, hallucinations and sometimes hydrophobia, which progress to delirium, coma and death. Around 17,400 people died from rabies in 2015, mainly in Asia and Africa.

Rabies is mentioned in the Codex of Eshnunna of around 1930 BC. The first vaccine was developed in 1885 by Louis Pasteur and Émile Roux. Prophylactic vaccination is used in people at high risk, pets and wild animals. Post-exposure prophylaxis, including vaccine and immunoglobulin, is completely effective if begun immediately after exposure, but survival is extremely rare once symptoms have begun.

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Zika fever is caused by the Zika virus, a mosquito-borne flavivirus first isolated in Uganda in 1947. Human infection is usually asymptomatic; a minority of cases have mild symptoms, which generally last 2–7 days and can include fever, conjunctivitis, joint pain, headache and a maculopapular rash. Infections in adults have occasionally been associated with Guillain–Barré syndrome. The incubation period is probably up to a week. The natural reservoir is unknown. The virus is transmitted by Aedes mosquitoes, mainly Aedes aegypti in tropical regions. During pregnancy, mother-to-child transmission can cause microcephaly (pictured) and other brain malformations in some babies. In one study, major abnormalities were seen in 42% of live births. The virus is present in semen and male-to-female sexual transmission has been documented; it is also found in breast milk and blood.

The first documented human outbreak occurred in 2007 in the Federated States of Micronesia. The virus is thought to have entered the Americas in around 2013, and an outbreak started in Brazil in 2015, spreading across the Americas and to Pacific, Asia and Africa, and leading the World Health Organization to consider Zika a Public Health Emergency of International Concern in February–November 2016. No specific treatment nor vaccine has been approved. Prevention involves mosquito control and condom use; women in areas where Zika was circulating were recommended to consider delaying pregnancy, and pregnant women were advised to avoid travel to affected areas.

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Myxomatosis is a disease of rabbits caused by Myxoma virus, a poxvirus in the genus Leporipoxvirus. The natural hosts are brush rabbits (Sylvilagus bachmani) in North America and tapeti (S. brasiliensis) in South and Central America, in which the myxoma virus causes only a mild disease, involving skin nodules. In European rabbits (Oryctolagus cuniculus), it causes a severe, often fatal, disease. Symptoms include fever, swelling of the eyelids and anogenital area, a mucopurulent ocular and nasal discharge, respiratory distress and hypothermia. Death generally occurs 10–12 days after infection. Myxoma virus is transmitted passively (without replication) by arthropod vectors, usually via the bites of mosquitoes and fleas, and also mites, flies and lice. It can also be transmitted by direct contact, and is shed in the ocular and nasal discharge and from eroded skin.

Myxoma virus was intentionally introduced in Australia, France and Chile in the 1950s to control wild European rabbit populations. This resulted in short-term 10–100-fold reductions in the rabbit population, followed by its recovery with the emergence of myxomatosis-resistant animals and attenuated virus variants. The introduction of myxomatosis is regarded as a classical example of host–pathogen coevolution following cross-species transmission of a pathogen to a naive host.

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