herpesviruses

ヘルペスウイルス科、ヘルペスウイルス

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Herpesviridae
Virus classification
Group:	Group I (dsDNA)
Order:	Herpesvirales
Family:	Herpesviridae
Genera
Subfamily: Alphaherpesvirinae Iltovirus Mardivirus Simplexvirus Varicellovirus Subfamily: Betaherpesvirinae Cytomegalovirus Muromegalovirus Proboscivirus Roseolovirus Subfamily: Gammaherpesvirinae Lymphocryptovirus Macavirus Percavirus Rhadinovirus

Herpesviridae is a large family of DNA viruses that cause diseases in animals, including humans. The members of this family are also known as herpesviruses. The family name is derived from the Greek word herpein ("to creep"), referring to the latent, recurring infections typical of this group of viruses. Herpesviridae can cause latent or lytic infections.

At least five species of Herpesviridae – HSV-1 and HSV-2 (both of which can cause orolabial herpes and genital herpes), Varicella zoster virus (which causes chicken-pox and shingles), Epstein-Barr virus (which causes mononucleosis), and Cytomegalovirus – are extremely widespread among humans. More than 90% of adults have been infected with at least one of these, and a latent form of the virus remains in most people. In total, there are 8 herpesvirus types that infect humans: herpes simplex viruses 1 and 2, varicella-zoster virus, EBV (Epstein-Barr virus), human cytomegalovirus, human herpesvirus 6, human herpesvirus 7, and Kaposi's sarcoma-associated herpesvirus. There are more than 130 herpesviruses, and some are from mammals, birds, fish, reptiles, amphibians, and molluscs.

Viral structure

Herpesviruses all share a common structure—all herpesviruses are composed of relatively large double-stranded, linear DNA genomes encoding 100-200 genes encased within an icosahedral protein cage called the capsid which is itself wrapped in a protein layer called the tegument containing both viral proteins and viral mRNAs and a lipid bilayer membrane called the envelope. This whole particle is known as a virion.

Genus	Structure	Symmetry	Capsid	Genomic Arrangement	Genomic Segmentation
Iltovirus	Spherical Pleomorphic	T=16	Enveloped	Linear	Monopartite
Proboscivirus	Spherical Pleomorphic	T=16	Enveloped	Linear	Monopartite
Cytomegalovirus	Spherical Pleomorphic	T=16	Enveloped	Linear	Monopartite
Mardivirus	Spherical Pleomorphic	T=16	Enveloped	Linear	Monopartite
Rhadinovirus	Spherical Pleomorphic	T=16	Enveloped	Linear	Monopartite
Macavirus	Spherical Pleomorphic	T=16	Enveloped	Linear	Monopartite
Roseolovirus	Spherical Pleomorphic	T=16	Enveloped	Linear	Monopartite
Simplexvirus	Spherical Pleomorphic	T=16	Enveloped	Linear	Monopartite
Scutavirus	Spherical Pleomorphic	T=16	Enveloped	Linear	Monopartite
Varicellovirus	Spherical Pleomorphic	T=16	Enveloped	Linear	Monopartite
Percavirus	Spherical Pleomorphic	T=16	Enveloped	Linear	Monopartite
Lymphocryptovirus	Spherical Pleomorphic	T=16	Enveloped	Linear	Monopartite
Muromegalovirus	Spherical Pleomorphic	T=16	Enveloped	Linear	Monopartite

Herpesvirus life cycle

All herpesviruses are nuclear-replicating—the viral DNA is transcribed to mRNA within the infected cell's nucleus.

Infection is initiated when a viral particle contacts a cell with specific types of receptor molecules on the cell surface. Following binding of viral envelope glycoproteins to cell membrane receptors, the virion is internalized and dismantled, allowing viral DNA to migrate to the cell nucleus. Within the nucleus, replication of viral DNA and transcription of viral genes occurs.

During symptomatic infection, infected cells transcribe lytic viral genes. In some host cells, a small number of viral genes termed latency associated transcript (LAT) accumulate instead. In this fashion the virus can persist in the cell (and thus the host) indefinitely. While primary infection is often accompanied by a self-limited period of clinical illness, long-term latency is symptom-free.

Reactivation of latent viruses has been implicated in a number of diseases (e.g. Shingles, Pityriasis Rosea). Following activation, transcription of viral genes transitions from latency-associated LAT to multiple lytic genes; these lead to enhanced replication and virus production. Often, lytic activation leads to cell death. Clinically, lytic activation is often accompanied by emergence of non-specific symptoms such as low grade fever, headache, sore throat, malaise, and rash as well as clinical signs such as swollen or tender lymph nodes and immunological findings such as reduced levels of natural killer cells.

Genus	Host Details	Tissue Tropism	Entry Details	Release Details	Replication Site	Assembly Site	Transmission
Iltovirus	Birds: galliform: psittacine	None	Cell receptor endocytosis	Budding	Nucleus	Nucleus	Oral-fecal; aerosol
Proboscivirus	Elephants	None	Glycoproteins	Budding	Nucleus	Nucleus	Contact
Cytomegalovirus	Humans; monkeys	Epithelial mucosa	Glycoproteins	Budding	Nucleus	Nucleus	Urine; saliva
Mardivirus	Chickens; turkeys; quail	None	Cell receptor endocytosis	Budding	Nucleus	Nucleus	Aerosol
Rhadinovirus	Humans; mammals	B-lymphocytes	Glycoproteins	Budding	Nucleus	Nucleus	Sex; saliva
Macavirus	Mammals	B-lymphocytes	Glycoproteins	Budding	Nucleus	Nucleus	Sex; saliva
Roseolovirus	Humans	T-cells; B-cells; NK-cell; monocytes; macrophages; epithelial	Glycoproteins	Budding	Nucleus	Nucleus	Respiratory contact
Simplexvirus	Humans; mammals	Epithelial mucosa	Cell receptor endocytosis	Budding	Nucleus	Nucleus	Saliva
Scutavirus	Sea turtles	None	Cell receptor endocytosis	Budding	Nucleus	Nucleus	Aerosol
Varicellovirus	Mammals	Epithelial mucosa	Glycoproteins	Budding	Nucleus	Nucleus	Aerosol
Percavirus	Mammals	B-lymphocytes	Glycoproteins	Budding	Nucleus	Nucleus	Sex; saliva
Lymphocryptovirus	Humans; mammals	B-lymphocytes	Glycoproteins	Budding	Nucleus	Nucleus	Saliva
Muromegalovirus	Rodents	Salivary glands	Glycoproteins	Budding	Nucleus	Nucleus	Contact

Taxonomy

Group: dsDNA

Order: Herpesvirales

• Family: Herpesviridae

  • Sub-Family: Alphaherpesvirinae

    • Genus: Iltovirus 

      • Gallid herpesvirus 1
      • Psittacid herpesvirus 1
    • Genus: Mardivirus 

      • Anatid herpesvirus 1
      • Columbid herpesvirus 1
      • Gallid herpesvirus 2
      • Gallid herpesvirus 3
      • Meleagrid herpesvirus 1
    • Genus: Scutavirus 

      • Chelonid herpesvirus 5
    • Genus: Simplexvirus 

      • Ateline herpesvirus 1
      • Bovine herpesvirus 2
      • Cercopithecine herpesvirus 2
      • Human herpesvirus 1
      • Human herpesvirus 2
      • Leporid herpesvirus 4
      • Macacine herpesvirus 1
      • Macropodid herpesvirus 1
      • Macropodid herpesvirus 2
      • Papiine herpesvirus 2
      • Saimiriine herpesvirus 1
    • Genus: Unassigned 

      • Chelonid herpesvirus 6
    • Genus: Varicellovirus 

      • Bovine herpesvirus 1
      • Bovine herpesvirus 5
      • Bubaline herpesvirus 1
      • Canid herpesvirus 1
      • Caprine herpesvirus 1
      • Cercopithecine herpesvirus 9
      • Cervid herpesvirus 1
      • Cervid herpesvirus 2
      • Equid herpesvirus 1
      • Equid herpesvirus 3
      • Equid herpesvirus 4
      • Equid herpesvirus 8
      • Equid herpesvirus 9
      • Felid herpesvirus 1
      • Human herpesvirus 3
      • Phocid herpesvirus 1
      • Suid herpesvirus 1
  • Sub-Family: Betaherpesvirinae

    • Genus: Cytomegalovirus 

      • Aotine herpesvirus 1
      • Cebine herpesvirus 1
      • Cercopithecine herpesvirus 5
      • Human herpesvirus 5
      • Macacine herpesvirus 3
      • Panine herpesvirus 2
      • Papiine herpesvirus 3
      • Saimiriine herpesvirus 4
    • Genus: Muromegalovirus 

      • Murid herpesvirus 1
      • Murid herpesvirus 2
      • Murid herpesvirus 8
    • Genus: Proboscivirus 

      • Elephantid herpesvirus 1
    • Genus: Roseolovirus 

      • Human herpesvirus 7
      • Human herpesvirus 6A
      • Human herpesvirus 6B
    • Genus: Unassigned 

      • Caviid herpesvirus 2
      • Suid herpesvirus 2
      • Tupaiid herpesvirus 1
  • Sub-Family: Gammaherpesvirinae

    • Genus: Lymphocryptovirus 

      • Callitrichine herpesvirus 3
      • Cercopithecine herpesvirus 14
      • Gorilline herpesvirus 1
      • Human herpesvirus 4
      • Macacine herpesvirus 4
      • Panine herpesvirus 1
      • Papiine herpesvirus 1
      • Pongine herpesvirus 2
    • Genus: Macavirus 

      • Alcelaphine herpesvirus 1
      • Alcelaphine herpesvirus 2
      • Bovine herpesvirus 6
      • Caprine herpesvirus 2
      • Hippotragine herpesvirus 1
      • Ovine herpesvirus 2
      • Suid herpesvirus 3
      • Suid herpesvirus 4
      • Suid herpesvirus 5
    • Genus: Percavirus 

      • Equid herpesvirus 2
      • Equid herpesvirus 5
      • Mustelid herpesvirus 1
    • Genus: Rhadinovirus 

      • Ateline herpesvirus 2
      • Ateline herpesvirus 3
      • Bovine herpesvirus 4
      • Cricetid herpesvirus 2
      • Human herpesvirus 8
      • Macacine herpesvirus 5
      • Murid herpesvirus 4
      • Murid herpesvirus 7
      • Saimiriine herpesvirus 2
    • Genus: Unassigned 

      • Equid herpesvirus 7
      • Phocid herpesvirus 2
      • Saguinine herpesvirus 1

The herpesvirus was first isolated from the blue wildebeest in 1960 by veterinary scientist Walter Plowright. The genus Herpesvirus was established in 1971 in the first report of the International Committee on Taxonomy of Viruses (ICTV). This genus consisted of 23 viruses and 4 groups of viruses. In the second ICTV report in 1976 this genus was elevated to family level - the Herpetoviridae. Because of possible confusion with viruses derived from reptiles this name was changed in the third report in 1979 to Herpesviridae. In this report the family Herpesviridae was divided into 3 subfamilies (Alphaherpesvirinae, Betaherpesvirinae and Gammaherpesvirinae) and 5 unnamed genera: 21 viruses were listed. In 2009 the family Herpesviridae was elevated to the order Herpesvirales. This elevation was necessitated by the discovery that the herpes viruses of fish and molluscs were only distantly related to those of birds and mammals. Two new families were created - the family Alloherpesviridae which incorporates bony fish and frog viruses and the family Malacoherpesviridae which contains those of molluscs.

This order currently has 3 families, 3 subfamilies plus 1 unassigned, 17 genera, 90 species and plus 48 as yet unassigned viruses.

Virus naming system

The system of naming herpes viruses was originated in 1973 and has been elaborated considerably since. The recommended naming system specified that each herpes virus should be named after the taxon (family or subfamily) to which its primary natural host belongs. The subfamily name is used for viruses from members of the family Bovidae or from primates (the virus name ending in –ine, e.g. bovine) and the host family name for other viruses (ending in –id, e.g. equid). Human herpes viruses have been treated as an exception (human rather than hominid). Following the host-derived term, the word herpes virus is added, followed by an Arabic numeral (1,2,3,...). These last two additions bear no implied meaning about taxonomic or biological properties of the virus.

Some exceptions to this system exist. A number of viruses' names (e.g. Epstein–Barr virus) are so widely used that it is impractical to attempt to insist on their replacement. This has led to a dual nomenclature in the literature for some herpes viruses. All herpes viruses described since this system was adopted have been named in accordance with it.

Evolution

The three mammalian subfamilies - Alpha, Beta and Gamma - arose approximately 180 million years ago to 220 million years ago. The major sublineages within these subfamilies were probably generated before the mammalian radiation of 80 million years ago to 60 million years ago. Speciations within sublineages took place in the last 80 million years probably with a major component of cospeciation with host lineages.

All the currently known bird and reptile species are alphaherpesviruses. Although the branching order of the herpes viruses has not yet been resolved, because herpes viruses and their hosts tend to coevolve this is suggestive that the alphaviruses may have been the earliest branch.

The date of evolution of the iltovirus genus has been estimated to be 200 million years ago while those of the mardivirus and simplex genera have been estimated to be between 150 million years ago and 100 million years ago.

Immune system evasions

Herpesviruses are known for their ability to establish lifelong infections. One way this is possible is through immune evasion. Herpesviruses have many different ways of evading the immune system. One such way is by encoding a protein mimicking human interleukin 10 (hIL-10) and another is by downregulation of the major histocompatibility complex II (MHC II) in infected cells.

cmvIL-10

Research conducted on cytomegalovirus (CMV) indicates that the viral human IL-10 homolog, cmvIL-10, is important in inhibiting pro-inflammatory cytokine synthesis. The cmvIL-10 protein has 27% identity with hIL-10 and only one conserved residue out of the nine amino acids that make up the functional site for cytokine synthesis inhibition on hIL-10. There is, however, much similarity in the functions of hIL-10 and cmvIL-10. Both have been shown to down regulate IFN-γ, IL-1α, GM-CSF, IL-6 and TNF-α, which are all pro-inflammatory cytokines. They have also been shown to play a role in downregulating MHC I and MHC II and up regulating HLA-G (non-classical MHC I). These two events allow for immune evasion by suppressing the cell-mediated immune response and natural killer cell response, respectively. The similarities between hIL-10 and cmvIL-10 may be explained by the fact that hIL-10 and cmvIL-10 both use the same cell surface receptor, the hIL-10 receptor. One difference in the function of hIL-10 and cmvIL-10 is that hIL-10 causes human peripheral blood mononuclear cells (PBMC) to both increase and decrease in proliferation whereas cmvIL-10 only causes a decrease in proliferation of PBMCs. This indicates that cmvIL-10 may lack the stimulatory effects that hIL-10 has on these cells.

It was found that cmvIL-10 functions through phosphorylation of the Stat3 protein. It was originally thought that this phosphorylation was a result of the JAK-STAT pathway. However, despite evidence that JAK does indeed phosphorylate Stat3, its inhibition has no significant influence on cytokine synthesis inhibition. Another protein, PI3K, was also found to phosphorylate Stat3. PI3K inhibition, unlike JAK inhibition, did have a significant impact on cytokine synthesis. The difference between PI3K and JAK in Stat3 phosphorylation is that PI3K phosphorylates Stat3 on the S727 residue whereas JAK phosphorylates Stat3 on the Y705 residue. This difference in phosphorylation positions seems to be the key factor in Stat3 activation leading to inhibition of pro-inflammatory cytokine synthesis. In fact, when a PI3K inhibitor is added to cells, the cytokine synthesis levels are significantly restored. The fact that cytokine levels are not completely restored indicates there is another pathway activated by cmvIL-10 that is inhibiting cytokine system nthesis. The proposed mechanism is that cmvIL-10 activates PI3K which in turn activates PKB (Akt). PKB may then activate mTOR, which may target Stat3 for phosphorylation on the S727 residue.

MHC downregulation

Another one of the many ways in which herpes viruses evade the immune system is by down regulation of MHC I and MHC II. This is observed in almost every human herpesvirus. Down regulation of MHC I and MHC II can come about by many different mechanisms, most causing the MHC to be absent from the cell surface. As discussed above, one way is by a viral chemokine homolog such as IL-10. Another mechanism to down regulate MHCs is to encode viral proteins that detain the newly formed MHC in the endoplasmic reticulum (ER). The MHC cannot reach the cell surface and therefore cannot activate the T cell response. The MHCs can also be targeted for destruction in the proteasome or lysosome. The ER protein TAP also plays a role in MHC down regulation. Viral proteins inhibit TAP preventing the MHC from picking up a viral antigen peptide. This prevents proper folding of the MHC and therefore the MHC does not reach the cell surface.

It is important to note that HLA-G is often up regulated in addition to downregulation of MHC I and MHC II. This prevents the natural killer cell response.

Human herpesvirus types

There are eight distinct viruses in this family known to cause disease in humans.

Human Herpesvirus (HHV) classification
Name	Synonym	Subfamily	Primary Target Cell	Pathophysiology	Site of Latency	Means of Spread
HHV‑1	Herpes simplex virus-1 (HSV-1)	α (Alpha)	Mucoepithelial	Oral and/or genital herpes (predominantly orofacial), as well as other herpes simplex infections	Neuron	Close contact (oral or sexually transmitted infection)
HHV-2	Herpes simplex virus-2 (HSV-2)	α	Mucoepithelial	Oral and/or genital herpes (predominantly genital), as well as other herpes simplex infections	Neuron	Close contact (oral or sexually transmitted disease)
HHV-3	Varicella zoster virus (VZV)	α	Mucoepithelial	Chickenpox and shingles	Neuron	Respiratory and close contact (including sexually transmitted disease)
HHV-4	Epstein-Barr virus (EBV), lymphocryptovirus	γ (Gamma)	B cells and epithelial cells	Infectious mononucleosis, Burkitt's lymphoma, CNS lymphoma in AIDS patients, post-transplant lymphoproliferative syndrome (PTLD), nasopharyngeal carcinoma, HIV-associated hairy leukoplakia	B cell	Close contact, transfusions, tissue transplant, and congenital
HHV-5	Cytomegalovirus (CMV)	β (Beta)	Monocyte, lymphocyte, and epithelial cells	Infectious mononucleosis-like syndrome, retinitis	Monocyte, lymphocyte, and ?	Saliva, urine, breast milk
HHV-6A and 6B	Roseolovirus, Herpes lymphotropic virus	β	T cells and ?	Sixth disease (roseola infantum or exanthem subitum)	T cells and ?	Respiratory and close contact?
HHV-7		β	T cells and ?	(roseola infantum or exanthem subitum)	T cells and ?	?
HHV-8	Kaposi's sarcoma-associated herpesvirus (KSHV), a type of rhadinovirus	γ	Lymphocyte and other cells	Kaposi's sarcoma, primary effusion lymphoma, some types of multicentric Castleman's disease	B cell	Close contact (sexual), saliva?

Herpesviruses of other animals

In addition to the herpesviruses considered endemic in humans, some viruses associated primarily with animals may infect humans. These are zoonotic infections:

Zoonotic Herpesviruses
Species	Type	Synonym	Subfamily	Human Pathophysiology
Macaque monkey	CeHV-1	Cercopithecine herpesvirus-1, (Monkey B virus)	α	Very unusual, with only approximately 25 human cases reported. Untreated infection is often deadly; sixteen of the 25 cases resulted in fatal encephalomyelitis. At least four cases resulted in survival with severe neurologic impairment. Symptom awareness and early treatment are important for laboratory workers facing exposure.
Mouse	MuHV‑4	Murid herpesvirus 68 (MHV-68)	γ	Zoonotic infection found in 4.5% of general population and more common in laboratory workers handling infected mice. ELISA tests show factor-of-four (x4) false positive results, due to antibody cross-reaction with other Herpes viruses.

Animal herpesviruses

In animal virology, the most important herpesviruses belong to the subfamily Alphaherpesvirinae. Research on pseudorabies virus (PrV), the causative agent of Aujeszky's disease in pigs, has pioneered animal disease control with genetically modified vaccines. PrV is now extensively studied as a model for basic processes during lytic herpesvirus infection, and for unraveling molecular mechanisms of herpesvirus neurotropism, whereas bovine herpesvirus 1, the causative agent of bovine infectious rhinotracheitis and pustular vulvovaginitis, is analyzed to elucidate molecular mechanisms of latency. The avian infectious laryngotracheitis virus is phylogenetically distant from these two viruses and serves to underline similarity and diversity within the Alphaherpesvirinae.

• Subfamily Alphaherpesvirinae

• • Genus Simplexvirus
    • Ateline herpesvirus 1, Spider monkey herpesvirus.
    • Bovine herpesvirus 2 causes bovine mammillitis and pseudo-lumpyskin disease.
    • Cercopithecine herpesvirus 1, also known as Herpes B virus, causes a Herpes simplex-like disease in Macaques, usually fatal if symptomatic and untreated in humans.
    • Fruit bat alphaherpesvirus 1
    • Leporid herpesvirus 4
    • Macacine herpesvirus 1
    • Macropodid herpesvirus 2
    • Papiine herpesvirus 2

• • Genus Varicellovirus
    • Bovine herpesvirus 1 causes infectious bovine rhinotracheitis, vaginitis, balanoposthitis, and abortion in cattle.
    • Bovine herpesvirus 5 causes encephalitis in cattle.
    • Bubaline herpesvirus 1
    • Caprine herpesvirus 1 causes conjunctivitis and respiratory disease in goats.
    • Canine herpesvirus 1 causes a severe hemorrhagic disease in puppies.
    • Cercopithecine herpesvirus 9
    • Cervid herpesvirus 1
    • Cervid herpesvirus 2
    • Elk herpesvirus 1
    • Equine herpesvirus 1 causes respiratory disease, neurological disease/paralysis, and spontaneous abortion in horses.
    • Equine herpesvirus 3 causes coital exanthema in horses.
    • Equine herpesvirus 4 causes rhinopneumonitis in horses.
    • Equine herpesvirus 8
    • Equine herpesvirus 9
    • Feline herpesvirus 1 causes feline viral rhinotracheitis and keratitis in cats.
    • Suid herpesvirus 1 causes Aujeszky's disease, also called pseudorabies.

• • Genus Mardivirus
    • Anatid herpesvirus 1
    • Columbiform herpesvirus 1
    • Gallid herpesvirus 2 causes Marek's disease.
    • Gallid herpesvirus 3 (GaHV-3 or MDV-2)
    • Meleagrid herpesvirus 1 (HVT)
    • Peacock herpesvirus 1

• • Genus Iltovirus
    • Gallid herpesvirus 1 causes infectious laryngotracheitis in birds.
    • Psittacid herpesvirus 1

Reptilian alphaherpesviruses

• Caretta caretta herpesvirus

• Chelonid herpesvirus 1
• Chelonid herpesvirus 2
• Chelonid herpesvirus 3
• Chelonid herpesvirus 4

• Chelonia mydas herpesvirus

• Coober herpesvirus

• Emydid herpesvirus 1
• Emydid herpesvirus 2

• Fibropapilloma associated herpes virus

• Gerrhosaurid herpesvirus 1
• Gerrhosaurid herpesvirus 2
• Gerrhosaurid herpesvirus 3

• Glyptemis herpesvirus 1
• Glyptemys herpesvirus 2

• Iguanid herpesvirus 1
• Iguanid herpesvirus 2

• Loggerhead orocutaneous herpesvirus
• Lung-eye-trachea associated herpesvirus

• Pelomedusid herpesvirus 1

• Red eared slider herpes virus

• Terrapene herpesvirus 1
• Terrapene herpesvirus 2

• Testudinid herpesvirus 1
• Testudinid herpesvirus 2
• Testudinid herpesvirus 3
• Testudinid herpesvirus 4

• Varanid herpesvirus 1

• Subfamily Betaherpesvirinae

• • Genus Proboscivirus

• • Porcine herpesvirus 2 causes inclusion body rhinitis in swine.

• Subfamily Gammaherpesvirinae

• • Genus Macavirus

• • Genus Percavirus

• • Genus Rhadinovirus
    • Alcelaphine herpesvirus 1 causes bovine malignant catarrhal fever.
    • Alcelaphine herpesvirus 2 causes an antelope and hartebeest version of MCF
    • Ateline herpesvirus 2
    • Bovine herpesvirus 4
    • Cercopithecine herpesvirus 17
    • Equine herpesvirus 2 causes equine cytomegalovirus infection.
    • Equine herpesvirus 5
    • Equine herpesvirus 7
    • Japanese macaque rhadinovirus
    • Leporid herpesvirus 1
    • Murid herpesvirus 4 Also known as Murine gammaherpesvirus-68 (MHV-68)

Research

Research is currently ongoing into a variety of side-effect or co-conditions related to the herpesviruses. These include:

• Alzheimer's disease
• atherosclerosis
• cholangiocarcinoma
• Crohn's disease
• chronic fatigue syndrome
• dysautonomia
• fibromyalgia
• Irritable bowel syndrome
• multiple sclerosis
• labile hypertension
• lupus
• pancreatic cancer
• pancreatitis
• pityriasis rosea
• Type II Diabetes
• Ménière's disease

References

External links

• ICTV International Committee on Taxonomy of Viruses (official site)
• Viralzone: Herpesviridae
• Animal viruses
• Article on Cercopithecine herpesvirus
• National B Virus Resource Center
• Pityriasis Rosea overview
• Herpes simplex: Host viral protein interactions.A database of Host/HSV-1 interactions
• Virus Pathogen Database and Analysis Resource (ViPR): Herpesviridae

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• Zhang X, Dawson CW, He Z, Huang P.SourceDepartment of Pathophysiology, Guangdong Medical College, Songshan Lake Industrial & Technology Park, Dongguan, Guangdong, China. zhangxn_2006@126.com.
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• Two human gamma-herpesviruses, Epstein-Barr virus and Kaposi's sarcoma associated herpesvirus/human herpesvirus 8 display oncogenic potential, causing benign and malignant lymphoproliferative disorders in genetically susceptible or immunosuppressed individuals. As a family of viruses that establish
• PMID 22170548

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• 造血細胞移植後HHV-6関連疾患:―最新の話題―

• 臨床血液 57(3), 298-306, 2016
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