Herpesviruses: overview of systematics, genomic complexity and life cycle

Aurélie Dotto-Maurel; Isabelle Arzul; Benjamin Morga; Germain Chevignon

doi:10.1186/s12985-025-02779-7

. 2025 May 22;22:155. doi: 10.1186/s12985-025-02779-7

Herpesviruses: overview of systematics, genomic complexity and life cycle

Aurélie Dotto-Maurel ^1,^✉, Isabelle Arzul ¹, Benjamin Morga ¹, Germain Chevignon ^1,^✉

PMCID: PMC12096621 PMID: 40399963

Abstract

Herpesviruses are double-stranded DNA viruses with distinct morphological features and are among the largest and most complex viruses. According to the International Committee on Taxonomy of Viruses (ICTV), in 2022, there were 133 herpesviruses classified into three families: Orthoherpesviridae, infecting mammals and birds; Malacoherpesviridae infecting marine molluscs; and Alloherpesviridae infecting fish and amphibians. Herpesviruses have a complex genomic architecture, characterised by unique regions flanked by repeated and inverted sequences. Unique regions can undergo rearrangements leading to the formation of genomic isomers, which could have important implications for the life cycle of the virus. Herpesviruses life cycle consists of two main phases: the lytic phase, during which viral genes are expressed and translated into viral proteins that regulate DNA replication, capsid formation and the production of new particles; and the persistence phase, in which the virus persists in the host without being eliminated by the immune system. This review offers an updated and comprehensive overview of the Herpesvirales order, detailing their morphological characteristics, providing an in-depth taxonomic classification, examining their genomic architecture and isomers, and describing their life cycle.

Keywords: Herpesvirus, Life cycle, Genomic architecture, Phylogeny

Background

Viruses play a major role in shaping life and driving evolution. They are the smallest and most abundant form of life, with an estimated 10³¹ particles in the biosphere, and occupy almost every ecosystem infecting all types of life forms [1–6]. Currently, there are four types of viral genome: double-stranded RNA (dsRNA), single-stranded RNA (ssRNA), double-stranded DNA (dsDNA), single-stranded DNA (ssDNA) (https://ictv.global/virus-properties, consulted on 28/01/25). Herpesviruses are dsDNA viruses belonging to the order Herpesvirales within the realm Duplodnaviria (Fig. 1) [7].

Fig. 1 — Overall representation of DNA viruses. Realms, kingdoms, phyla, classes and orders are shown for all viruses whereas families, subfamilies are shown for herpesviruses (adapted from https://ictv.global/taxonomy/visual-browser)

Herpesviruses are ubiquitous throughout the world and have co-existed with mankind for as long as there have been written records. Interestingly, ancient Egyptian hieroglyphics document the presence of herpetic lesions, suggesting that these viruses have been a concern issue for millennia [8]. The origin of herpesviruses has been studied since the 1990s, with several estimates suggesting that their most recent common ancestor existed between 150.1 and 209.9 million years ago, dating back to the Jurassic/Triassic periods [9–12]. It has long been thought that herpesviruses evolve with their hosts primarily through co-speciation [10, 13–15]. However, as the amount of genomic sequence available has increased and bioinformatic tools have evolved, the explanation for herpesviruses evolution has shifted. Current evidence suggest that host-switching events and intra-host speciation play a more important role in their evolutionary history [9, 13, 16].

Historically, the classification of herpesviruses has been defined on the basis of virion architecture and host species [17]. In recent decades, herpesvirus genetic data have accumulated and become extensive, with more than 1000 complete genome sequences now available in public databases (https://www.ncbi.nlm.nih.gov/datasets/genome/?taxon=548681, consulted on 28/01/2025). Currently, and based on genetic content, 133 distinct herpesvirus species have been identified and grouped into three families: Orthoherpesviridae, Malacoherpesviridae and Alloherpesviridae [7, 18] (https://ictv.global/taxonomy, consulted on 28/01/25). In addition, four new species (Babylonia areolata herpesvirus [19], Bufonid herpesvirus 1 [20], Lake sturgeon herpesvirus 1 [21] and Silurid herpesvirus 1 [22] have not yet been classified.

Herpesviruses are capable of infecting a wide range of animals including mammals, birds, fish, and invertebrates (marine molluscs) [13, 23]. Despite some differences in host specificity, tissue tropism, replication kinetics and pathogenic potential, herpesviruses share a common virion morphology and life cycle.

Most available reviews focus on the family Orthoherpesviridae, and more specifically the eight human herpesviruses species [8, 24–26]. The last comprehensive review providing a global overview of herpesvirus systematics, genomic architectures and lifecycles was published in 2013, when the order Herpesvirales comprised 90 herpesvirus species [27, 28]. Since then, 43 additional herpesviruses have been characterised and included in the ICTV classification while 4 new species need to be included.

Therefore, this review seeks to provide an updated and comprehensive overview of the order Herpesvirales with the aims to detail their morphological features, offer a complete and up to date taxonomic classification, and present an in-depth analysis of their genomic architecture. Additionally, the review includes a general description of their life cycle. This synthesis is intended to serve as a valuable reference for researchers, facilitating a deeper understanding of the biology, evolution, and significance of these viruses.

Herpesviruses virion structure

Herpesviruses, members of the order Herpesvirales are double-stranded DNA (dsDNA) viruses with morphological characteristics that distinguish them from other viruses. The mature herpesvirus virion structure has been described for 27 of the 49 alphaherpesviruses, 17 of the 27 betaherpesviruses, 18 of the 42 gammaherpesviruses, 2 of the 3 malacoherpesviruses and 12 of the 16 alloherpesviruses and is composed of an envelope surrounding a tegument, a protein-rich matrix between the envelope and the capsid. The capsid encloses the core consisting in tightly packed dsDNA genome (Fig. 2). The size of herpesvirus mature virion varies between 120 to 260 nm in diameter, depending on the thickness of the tegument and the state of the envelope [8].

Fig. 2 — Schematic representation of herpesvirus virion

Envelope

The envelope is a lipid bilayer that maintains the quasi-spherical shape of the virion. Its trilaminar appearance has been demonstrated by electron microscopy [29]. The envelope consists mainly of virally encoded glycoproteins (Fig. 2). The number and relative amounts of viral glycoproteins vary among herpesviruses. For example, in Simplexvirus humanalpha 1, also known as Human Simplex Virus 1 (HSV-1), the envelope contains at least 11 different virion-associated glycoproteins and the copy number of individual glycoproteins can exceed 1,000 per virion [8]. The glycoproteins form numerous protrusions on the virion envelope that are more numerous and shorter than those found on the surface of many other enveloped viruses [8].

Tegument

The tegument is sometimes asymmetrically distributed and its thickness can vary depending on the location of the virion with the infected cell and is determined by the virus rather than the host [30]. The tegument contains more than 20 different virally encoded proteins, some of which are present in hundreds of copies per virion [8].

Capsid

The structural features of the capsid are characteristic of all herpesviruses. It measures 100 nm in diameter, contains 161 capsomers (150 hexons and 11 pentons) and has a capsid triangulation number of T = 16 [31–33]. Non-enveloped capsids are present in infected cells in three main forms: A-, B- and C-capsids, where A-capsids have no core structure and B-capsids contain the assembly scaffold but no genome. Only the C-capsids contain the complete dsDNA herpesvirus genome [8].

Core

The core of the mature virion contains the viral genome as a single molecule in the form of dsDNA packed in an ordered manner in the form of a torus [33–36]. The exact arrangement of the DNA in the torus is not known but the DNA is tightly packed so that the internal volume of the capsid is approximately equal to the cylindrical volume of the genome [8, 37].

Towards a better characterization of herpesvirus particle structure

In total there is less than 60% of the described herpesvirus species for which the virion structure has been characterised. Furthermore, the number of proteins contained in the virions is not precisely known and may vary from one virus to another. Proteomics analyses have identified up to 71 structure-related proteins in virions of human, mice and monkey herpesviruses. These proteins are distributed as follows: 4 to 7 in the nucleocapsid, 9 to 20 in the tegument and 4 to 19 in the envelope plus a number of proteins whose location within virions is unknown [8, 38–42]. Recent advances including high-resolution cryo-electron microscopy and integrative structural approaches provide powerful tools to better characterize herpesvirus particles. For example, cryo-electron-microscopy has revealed significant differences between A- and B-capsids and mature C-capsids, providing new opportunities to understand why only the C-capsids contain the complete dsDNA herpesvirus genome [43]. Moreover, native mass spectrometry could be used to accurately determine the mass of viral capsids and identify the proteins they contain [44]. Advancing the structural characterisation of herpesvirus virions would greatly improve our understanding of how these viruses infect their hosts and could be helpful to design not only new diagnostic tools but also new strategies to block infection.

Herpesvirales clade

Herpesviruses are host-specific pathogens classified into three families on the basis of the DNA polymerase sequences: Orthoherpesviridae infecting mammals, birds, and reptiles (118 species); Alloherpesviridae infecting fish and amphibians (16 species); and Malacoherpesviridae infecting marine molluscs (3 species) (Fig. 3A).

Fig. 3 — *Herpesvirales* phylogeny based on DNA polymerase sequences. DNA polymerase protein sequences from all available herpesviruses were extracted from public databases and aligned using Mafft v.7.520.A phylogenetic tree was constructed from the alignment file using phyml v.3.3.2 and the LG model with gamma distribution, invariant sites and amino acid frequencies. The resulting tree was then manually improved using Illustrator to add the names abbreviations, genomic architecture, size, isomers and latency characterization of the herpesviruses and the host they infect. A DNA polymerase tree, ICTV names, abbreviations and hosts. B Size and genomic architecture, grey colored sizes correspond to partial genome and black colored sizes correspond to complete genome. U: Unique region, TR: Terminal Repeat, IR: Internal Repeat, UL: Unique Long, US: Unique Short, TRL/IRL: Terminal/Inverted Repeat Long, TRS/IRS: Terminal/Internal Repeat Short, X: X region, a: a region, n: number of the repeat. C Genomic isomers. D Ability to establish persistence described in literature. L: latency demonstrated, P: Persistence described.

Orthoherpesviridae

The family Orthoherpesviridae contains 118 described herpesviruses species and is divided into three subfamilies: Alphaherpesvirinae with 49 species; Betaherpesvirinae with 27 species; and Gammaherpesvirinae with 42 species (https://ictv.global/taxonomy, consulted on 28/01/25).

Alphaherpesvirinae

Viruses in this subfamily include the Simplexvirus genus infecting humans, primates, rabbits, marsupials and bats the Varicellovirus genus infecting humans, cattle, dogs, primates, bovids, deer, horses, donkeys, giraffes, felines, whales, seals and swine the Scutavirus genus infecting marine and terrestrial turtle, Mardivirus and Iltovirus genera both infecting birds (Fig. 3A) (Table 1).

Table 1.

Summary table of viruses of the subfamily Alphaherpervirinae

	Scientific name	Common name	Abbv	Acc. Number (ICTV)	Reference Acc. Number	Host	Complete genomes	Sequencing technology for ref. genome
Simplexvirus	atelinealpha1	Ateline alphaherpesvirus 1	HVA1	KY385637	[45]	Primate	1	Sanger, Illumina
	bovinealpha2	Bovine alphaherpesvirus 2	BMV	MT862163	[46]	Cattle	2	Sanger, Illumina, ONT
	cercopithecinealpha2	Cercopithecine alphaherpesvirus 2	SA8	AY714813	[47]	Primate	1	Sanger
	humanalpha1	Herpes simplex virus type 1	HSV1	JN555585	[48]	Human	71	Illumina
	humanalpha2	Herpes simplex virus type 2	HSV2	JN561323	[48]	Human	14	Illumina
	leporidalpha4	Leporid herpesvirus 4	LHV4	JQ596859	[49]	Lepus	1	454
	macacinealpha1	Macacine alphaherpesvirus 1	BV	AF533768	[50]	Primate	18	Sanger
	macacinealpha2	Macacine alphaherpesvirus 2	McAHV2	KY628968	[45]	Primate	1	Sanger, Illumina
	macacinealpha3	Macacine alphaherpesvirus 3	McAHV3	KY628970	[45]	Primate	1	Sanger, Illumina
	macropodidalpha1	Macropodid alphaherpesvirus 1	MaAHV1	KT594769	[51]	Marsupial	1	Illumina
	macropodidalpha2	Macropodid alphaherpesvirus 2	MaAHV2	MT900475	[52]	Marsupial	1	Illumina
	macropodidalpha4	Macropodid alphaherpesvirus 4	MaAHV4	MT900474	[52]	Marsupial	1	Illumina
	paninealpha3	Panine alphaherpesvirus 3	ChHV	JQ360576	[53]	Primate	1	454
	papiinealpha2	Herpesvirus papio 2	HPV2	DQ149153	[54]	Primate	6	Sanger
	pteropodidalpha1	Fruit bat alphaherpesvirus 1	FBAHV1	AB825953	[55]	Bat	1	454
	pteropodidalpha2	Bat alphaherpesvirus	PLAHV	LC492974	[56]	Bat	1	Sanger, ONT
	saimiriinealpha1	Herpesvirus saimiri 1	HVS1	HM625781	[57]	Primate	1	Sanger, 454
Mardivirus	anatidalpha1	Duck enteritis virus	DEV	JF999965	[58]	Bird	10	454
	columbidalpha1	Pigeon herpesvirus	PHV	KX589235	[59]	Bird	2	Sanger
	gallidalpha2	Marek's disease virus	MDV	AF243438	[60]	Bird	65	Sanger
	gallidalpha3	Gallid herpesvirus 3	GaAHV3	HQ840738	[61]	Bird	2	454
	meleagridalpha1	Turkey herpesvirus	HVT	AF291866	[62]	Bird	1	Sanger
	spheniscidalpha1	Spheniscid alphaherpesvirus 1	SpAHV1	LT608135	[63]	Bird	2	454
Varicellovirus	bovinealpha1	Infectious bovine rhinotracheitis virus	IBRV	JX898220	[64]	Cattle	50	Sanger, Illumina
	bovinealpha5	Bovine encephalitis herpesvirus	BoAHV5	AY261359	[65]	Cattle	4	Sanger
	bubalinealpha1	Water buffalo herpesvirus	BuAHV1	KU936049	[66]	Cattle	1	Illumina
	canidalpha1	Canine herpesvirus	CHV	KT819633	[67]	Dog	11	Illumina
	caprinealpha1	Goat herpesvirus	CpAHV1	MG989243	[68]	Bovid	1	Illumina
	cercopithecinealpha9	Simian varicella virus	SVV	AF275348	[69]	Primate	1	Sanger
	cervidalpha1	Cervid alphaherpesvirus 1	CvAHV1	MH036942	Unpublished	Deer	1	Illumina
	cervidalpha2	Cervid alphaherpesvirus 2	CvAHV2	MH036943	Unpublished	Deer	1	Illumina
	cervidalpha3	Cervid alphaherpesvirus 3	CvAHV3	MH036941	Unpublished	Deer	1	Illumina
	equidalpha1	Equine abortion virus	EAV	AY665713	[70]	Horse	28	Sanger
	equidalpha3	Equine coital exanthema virus	EqAHV3	KM051845	[71]	Horse	1	454, Illumina
	equidalpha4	Equine rhinopneumonitis virus	EqAHV4	AF030027	[72]	Horse	10	Sanger
	equidalpha6	Ulcerative stomatitis in donkeys	AsHV	MT012704	[73]	Donkey	1	Illumina
	equidalpha8	Asinine herpesvirus 3	EqAHV8	MF431611	[74]	Horse	6	Illumina
	equidalpha9	Equid alphaherpesvirus 9	EqAHV9	AP010838	[75]	Giraffe	1	Sanger
	felidalpha1	Feline viral rhinotracheitis virus	FVRV	FJ478159	[76]	Feline	37	454
	humanalpha3	Varicella-zoster virus	VZV	X04370	[77]	Human	21	Sanger
	monodontidalpha1	Beluga whale alphaherpesvirus 1	MoAHV1	MF678601	[78]	Whale	1	Illumina
	phocidalpha1	Phocid herpesvirus 1	PhHV1	MH509440	[79]	Seal	0	-
	suidalpha1	Pseudorabies virus	PRV	JF797218	[80]	Swine	51	Illumina
Iltovirus	cacatuidalpha2	Cacatuid herpesvirus 2	CaHV-2	MK360902	[81]	Bird	1	Illumina
	gallidalpha1	Infectious laryngotracheitis virus	ILTV	JN596962	[82]	Bird	65	454
	psittacidalpha1	Pacheco's disease virus	PDV	AY372243	[83]	Bird	1	Sanger
	psittacidalpha5	Psittacid alphaherpesvirus 5	PsHV-5	MK955929	[84]	Bird	2	Illumina
Scutavirus	chelonidalpha5	Chelonid alphaherpesvirus 5	ChHV5	HQ878327	[85]	Turtle	0	-
Scutavirus	testudinidalpha3	Testudinid herpesvirus 3	TeHV3	KM924292	[86]	Turtle	5	Illumna

Open in a new tab

This table contains the scientific and common names, abbreviations, accession numbers from ICTV, related articles, hosts, number of complete genomes available on NCBI and the sequencing technology used for the reference genome (according to the accession number and related articles) of viruses of the Alphaherpesvirinae subfamily

Betaherpesvirinae

Viruses in this subfamily include the Cytomegalovirus genus infecting primates and humans, the Muromegalovirus genus infecting mice and rats, the Proboscivirus genus infecting elephants, the Quwivirus genus infecting rodents, and bats, and the Roseolovirus genus infecting humans, primates, swine and mice (Fig. 3A) (Table 2).

Table 2.

Summary table of viruses of the subfamily Betaherpervirinae

	Scientific name	Common name	Abbv	Acc. Number (ICTV)	Reference Acc. Number	Host	Complete genomes	Sequencing technology for ref. genome
Cytomegalovirus	aotinebeta1	Aotine betaherpesvirus 1	MCMV	FJ483970	Unpublished	Primate	1	not published
	cebinebeta1	Cebine herpesvirus 1	CMCMV	JQ264772	Unpublished	Primate	0	-
	cercopithecinebeta5	Cercopithecine betaherpesvirus 5	SCMV	FJ483968	Unpublished	Primate	3	not published
	humanbeta5	Human cytomegalovirus	HCMV	AY446894	[87]	Human	345	Sanger
	macacinebeta3	Rhesus cytomegalovirus	RhCMV	AY186194	[88]	Primate	30	Sanger
	macacinebeta8	Macaque cytomegalovirus	McBHV8	JN227533	[89]	Primate	7	Illumina
	mandrillinbeta1	Mandrillus leucophaeus cytomegalovirus	MdBHV1	KR297253	[90]	Primate	1	454
	paninebeta2	Panine betaherpesvirus 2	CCMV	AF480884	[91]	Primate	1	Sanger
	papiinebeta3	Olive baboon cytomegalovirus	BCMV	AC090446	Unpublished	Primate	0	-
	papiinebeta4	Chacma baboon cytomegalovirus	PaBHV4	KR351281	[90]	Primate	1	454
	saimiriinebeta4	Saimiriine betaherpesvirus 4	SMCMV	FJ483967	Unpublished	Primate	1	not published
Muromegalovirus	muridbeta1	Murine cytomegalovirus	MCMV	GU305914	[92]	Mice	15	Sanger
	muridbeta2	Rat cytomegalovirus strain Maastricht	RCMV	AF232689	[93]	Rat	1	Sanger
	muridbeta8	Rat cytomegalovirus strain England	RCMVE	JX867617	[94]	Rat	3	454, Sanger
Quwivirus	caviidbeta2	Guinea pig cytomegalovirus	GPCMV	KC503762	[95]	Rodent	2	Illumina
	miniopteridbeta1	Miniopterus schreibersii herpesvirus	MsHV	JQ805139	[96]	Bat	1	454
	tupaiidbeta1	Tupaiid betaherpesvirus 1	THV	AF281817	[97]	Shrew	1	Sanger
Roseolovirus	humanbeta7	Human herpesvirus 7	HHV7	AF037218	[98]	Human	2	Sanger
	humanbeta6a	Human herpesvirus 6A	HHV6A	X83413	[99]	Human	4	Sanger
	humanbeta6b	Human herpesvirus 6B	HHV6B	AF157706	[100]	Human	3	Sanger
	macacinebeta9	Macaca nemestrina herpesvirus 7	MneHV7	KU351741	[101]	Primate	1	Illumina
	muridbeta3	Mouse thymic virus	MTV	KY355735	[102]	Mice	2	Illumina, PacBio, Sanger
	suidbeta2	Porcine cytomegalovirus	PCMV	KF017583	Unpublished	Swine	1	Illumina
Proboscivirus	elephantidbeta1	Elephantid endotheliotropic herpesvirus 1	EEHV1	KC462165	[103]	Elephant	5	Illumina
	elephantidbeta3	Elephantid endotheliotropic herpesvirus 3	EEHV3	MN373268	[104]	Elephant	0	-
	elephantidbeta4	Elephantid endotheliotropic herpesvirus 4	EEHV4	KT832477	[105]	Elephant	1	Sanger, Illumina
	elephantidbeta5	Elephantid endotheliotropic herpesvirus 5	EEHV5	KF921519	[103]	Elephant	2	Illumina

Open in a new tab

Gammaherpesvirinae

Viruses in this subfamily include the Lymphocryptovirus genus infecting humans and primates, the Bossavirus genus infecting dolphins, the Macavirus infecting bovids, cattle, swine and hipparion, the Manticavirus genus infecting marsupials, the Patagivirus genus infecting bats, the Percavirus genus infecting horses, felines, badgers, seals and bats and the Rhadinovirus genus infecting primates, cattle, rodents, mice and humans (Fig. 3A) (Table 3).

Table 3.

Summary table of viruses of the subfamily Gammaherpervirinae

	Scientific name	Common name	Abbv	Acc. Number (ICTV)	Reference Acc. Number	Host	Complete genomes	Sequencing technology for ref. genome
Bossavirus	delphinidgamma1	Common bottlenose dolphin gammaherpesvirus 1	DeGHV1	KY965444	[106]	Dolphin	1	Illumina
Macavirus	alcelaphinegamma1	Wildebeest-associated malignant catarrhal fever virus	MCFV	AF005370	[107]	Bovid	0	-
	alcelaphinegamma2	Hartebeest-associated malignant catarrhal fever virus	AlGHV2	KF274499	[108]	Bovid	1	Sanger
	bovinegamma6	Bovine lymphotropic herpesvirus	BLV	KJ705001	[109]	Cattle	1	Illumina
	caprinegamma2	Caprine herpesvirus 2	CpHV2	AF283477	[110]	Bovid	0	-
	hippotraginegamma1	Roan antelope herpesvirus	HiGHV1	DQ083950	[111]	Hipparion	0	-
	ovinegamma2	Sheep-associated malignant catarrhal fever virus	OvGHV2	AY839756	[112]	Bovid	2	Sanger
	suidgamma3	Porcine lymphotropic herpesvirus 1	PLVH1	AF478169	[113]	Swine	0	-
	suidgamma4	Porcine lymphotropic herpesvirus 2	PLHV2	AY170317	[114]	Swine	0	-
	suidgamma5	Porcine lymphotropic herpesvirus 3	PLHV3	AY170316	[114]	Swine	0	-
Percavirus	equidgamma2	Equine herpesvirus 2	EHV2	U20824	[115]	Horse	20	Sanger
	equidgamma5	Equine herpesvirus 5	EHV5	KM924295	[116]	Horse	2	Illumina
	felidgamma1	Felis catus gammaherpesvirus 1	FcaGHV1	KT595939	[117]	Feline	1	Sanger, Illumina
	mustelidgamma1	Badger herpesvirus	BadHV	AF376034	[118]	Badger	0	-
	phocidgamma3	Harp seal herpesvirus	HaSHV	KP136799	[119]	Seal	0	-
	rhinolophidgamma1	Rhinolophus gammaherpesvirus 1	RGHV-1	LC333428	[120]	Bat	1	Ion Torrent
	vespertilionidgamma1	Bat gammaherpesvirus 8	VeGHV1	KU220026	[121]	Bat	0	-
Rhadinovirus	atelinegamma2	Herpesvirus ateles strain 810	AtGHV2	M22036	[122]	Primate	0	-
	atelinegamma3	Herpesvirus ateles	HVA	AF083424	[123]	Primate	1	Sanger
	bovinegamma4	Bovine herpesvirus 4	BHV4	AF318573	[124]	Cattle	0	-
	colobinegamma1	Colobine gammaherpesvirus 1	CbGHV1	MH932584	[125]	Primate	1	Illumina
	cricetidgamma2	Rodent herpesvirus Peru	RHVP	HQ221963	[126]	Rodent	1	454
	humangamma8	Kaposi's sarcoma-associated herpesvirus	KSHV	AF148805	[127]	Human	37	Sanger
	macacinegamma11	Japanese macaque rhadinovirus	JMRV	AY528864	[128]	Primate	3	Sanger
	macacinegamma12	Pig-tailed macaque rhadinovirus 2	MneRV2	KP265674	[129]	Primate	1	Sanger
	macacinegamma5	Rhesus rhadinovirus	RRV	AF083501	[130]	Primate	5	Sanger
	macacinegamma8	Retroperitoneal fibromatosis-associated herpesvirus	RFHV	KF703446	[131]	Primate	1	Sanger, Illumina
	muridgamma4	Murine gammaherpesvirus 68	MHV68	U97553	[132]	Mice	3	Sanger
	muridgamma7	Wood mouse herpesvirus	WMHV	GQ169129	[133]	Mice	1	Sanger
	saimiriinegamma2	Herpesvirus saimiri	HVS	X64346	[134]	Primate	1	Sanger
Patagivirus	vespertilionidgamma3	Eptesicus fuscus gammaherpesvirus	EfHV	MF385016	[135]	Bat	1	Illumina, ONT
Manticavirus	phascolarctidgamma1	Koala gammaherpesvirus 1	PcGHV1	MG452722	[136]	Marsupial	0	-
Manticavirus	vombatidgamma1	Wombat gammaherpesvirus 1	VoGHV1	MG452721	[136]	Marsupial	0	-
Lymphocryptovirus	callitrichinegamma3	Marmoset lymphosarcoma virus	ClGHV3	AF319782	[137]	Primate	1	Sanger
	gorillinegamma1	Pongine herpesvirus 3	GoGHV1	AJ581752	[138]	Primate	0	-
	humangamma4	Epstein-Barr virus	EBV	AJ507799	[139]	Human	311	Sanger
	macacinegamma10	cynomolgus macaque lymphocryptovirus	McGHV10	KP676001	[140]	Primate	1	Illumina
	macacinegamma13	Macaca arctoides gammaherpesvirus 1	McGHV13	NC_076963	[141]	Primate	1	Sanger, Illumina
	macacinegamma4	rhesus lymphocryptovirus	RLV	AY037858	[142]	Primate	1	Sanger
	paninegamma1	Pongine herpesvirus 1	PnGHV1	AJ581751	[138]	Primate	0	-
	papiinegamma1	Cercopithecine herpesvirus 12	PaGHV1	AJ581750	[138]	Primate	0	-
	ponginegamma2	Pongine herpesvirus 2	PoGHV2	AJ581753	[138]	Primate	0	-

Open in a new tab

Malacoherpesviridae

Only few species have been characterized in the Malacoherpesviridae family. This family encompasses viruses infecting marine molluscs such as oysters, abalones and sea snails. To date three viruses species are characterized in this family: the abalone herpesvirus Aurivirus haliotidmalaco1, also known as Abalone Herpesvirus (AbHV-1), the gastropod herpesvirus Babylonia areolata herpesvirus (BaHV) (not classified by ICTV yet) and the oyster herpesvirus Ostreavirus ostreidmalaco1, better known as Ostreid Herpesvirus Type 1 (OsHV-1) (Fig. 3A) (Table 4) (https://ictv.global/taxonomy, consulted on 28/01/2025).

Table 4.

Summary table of viruses of the family Malacoherpesviridae

Scientific name	Common name	Abbv	Acc. Number (ITCV)	Reference Acc. Number	Host	Complete genomes	Sequencing technology for ref. genome
Aurivirus haliotidmalaco1	Abalone herpesvirus	AbHV1	JX453331	[143]	Abalone	3	Sanger, 454, Illumina
Babylonia areolata herpesvirus	Gastropod herpesvirus	-	BK064993	[19]	Sea Snail	1	Illumina, ONT
Ostreavirus ostreidmalaco1	Ostreid herpesvirus type 1	OsHV1	AY509253	[32]	Oyster	28	Sanger

Open in a new tab

Alloherpesviridae

The viruses of the Alloherpesviridae family contains 13 characterised species, including 3 herpesviruses not classified yet by ICTV (Table 5, herpesviruses with an asterisk), clustered into four genera: the genus Batravirus infecting frogs, the genus Cyvirus infecting eels and carps, the genus Ictavirus infecting sturgeons and catfishes, and the genus Salmovirus infecting salmons (Fig. 3A) (Table 5) (https://ictv.global/taxonomy, consulted on 28/01/25).

Table 5.

Summary table of viruses of the family Alloherpesviridae

	Scientific name	Common name	Abbreviations	Acc. Number (ICTV)	Reference acc. Number	Host	Complete genomes	Sequencing technology for ref. genome
Batravirus	ranidallo1	Lucké tumor herpesvirus	LTHV	DQ665917	[144]	Frog	1	Sanger
	ranidallo2	Frog virus 4	FV4	DQ665652	[144]	Frog	1	Sanger
	ranidallo3	Ranid herpesvirus 3	RaHV3	KX832224	[145]	Frog	1	Illumina
	Bufonid herpesvirus 1		BfHV1	NC_040681	[20]	Frog	1	Illumina
Ictavirus	Lake sturgeon herpesvirus 1		LSHV	OK485036	[21]	Sturgeon	1	Illumina
	Silurid herpesvirus 1			MH048901	[22]	Catfish	1	Illumina
	acipenseridallo2	White sturgeon herpesvirus 2	AciHV2	FJ815289	[146]	Sturgeon	0	-
	ictaluridallo1	Channel catfish virus	IcHV1	M75136	[147]	Catfish	2	Sanger
	ictaluridallo2	Black bullhead herpesvirus	IcHV2	MG271984	[148]	Catfish	1	Sanger, Illumina
Cyvirus	anguillidallo1	Japanese eel herpesvirus	AngHV1	FJ940765	[149]	Eel	3	Sanger
	cyprinidallo1	Carp pox herpesvirus	CyHV1	JQ815363	[150]	Carp	1	Sanger
	cyprinidallo2	Goldfish hematopoietic necrosis virus	CyHV2	JQ815364	[150]	Carp	6	Illumina
	cyprinidallo3	Koi herpesvirus	KHV	DQ657948	[151]	Carp	21	Sanger
Salmovirus	salmonidallo1	Herpesvirus salmonis	SalHV1	AF023673	[152]	Salmon	0	-
	salmonidallo2	Oncorhyncus masou herpesvirus	SalHV2	EU349274	[153]	Salmon	0	-
	salmonidallo3	Epizootic epitheliotropic disease virus	SalHV3	EU349277	[153]	Salmon	0	-

Open in a new tab

Perspective on Herpesvirus classification

While the Orthoherpesviridae is the largest and most extensively described family, some studies suggest that the Alloherpesviridae and Malacoherpesviridae families may contain a greater number of as yet uncharacterised herpesviruses [154, 155]. Indeed, interrogation of publicly available sequencing data has recently led to the identification and characterisation of additional herpesviruses, as demonstrated by the detection of ancient HSV-1 and Simplexvirus humanalpha2 (HSV-2), strains in Neanderthal genome data [156], and the discovery of four novel malacoherpes-like viruses in SRA database samples [155]. In addition, environmental DNA sequencing-based approaches could facilitate detecting herpesviruses infecting hosts that are difficult to sample, such as terrestrial or marine wildlife species [157, 158]. Finally, methods such as the use of degenerated primers, loop-mediated isothermal amplification, microparticles, or tangential flow filtration, could further improve the detection and characterization of previously unknown herpesviruses [159–163]. While many herpesviruses remain uncharacterized, the 137 currently known herpesvirus species share a highly conserved virion structure,

consisting of a large, linear, double-stranded DNA genome, but exhibit considerable diversity in genetic content and genomic architecture.

Genomic architecture of Herpesvirales

Herpesvirus DNA genomes are linear, double stranded and their length varies from 119 kb for the Rhadinovirus muridgamma4 (MHV68) [132] to 295 kb for Cyvirus cyprinidallo3 (CyHV3) [151] (Fig. 3B).

An interesting feature of herpesviruses is their distinctive genomic organization [28]. In this section, the different genomic architectures of herpesviruses complete genome are described. To this end, genomic data were collected from the literature and from the genomic public database. For genome without known genomic structure, characterisation was performed manually by visualising genome via the self-alignment dotplot tool implemented in Geneious Prime (https://www.geneious.com/features/prime).

Seven distinct genomic architecture have been described:

Arch-1: consisting of two unique regions named Unique Long (U_L) and Unique Short (U_S) with the U_S region flanked by inverted repeat regions (IR_S: Internal Repeat Short and TR_S: Terminal Repeat Short), (i.e. U_L—IR_S—U_S—TR_S) (Table 6).
Arch-2: consisting of two unique regions (U_L and U_S) both flanked by inverted repeat sequences (IR_L: Internal Repeat Long, and TR_L: Terminal Repeat Long for the U_L and IR_S/TR_S for the U_S region), (i.e., TR_L—U_L—IR_L—IR_S—U_S—TR_S) (Table 6).
Arch-3: similar to Arch-2 with three copies of “a” region, located at both ends of the genome and between the IR_L and IR_S regions (i.e., a-TR_L—U_L—IR_L—a—IR_S—U_S—TR_S-a) (Table 6).
Arch-4: consisting of a unique DNA sequence (i.e. U) without any repeat sequences.
Arch-5: consisting in a unique long region (U) flanked by direct repeat regions (i.e. TR—U—TR) (Table 6).
Arch-6: consisting of terminal repeats (TR) located at either end of the genome, together with internal repeats (IR) interspersed throughout. Unlike the linked repeat arrangement seen in previous genomic architectures, these repeats are not directly linked in pairs. (i.e. TR—[U_(n)—IR_(n)]_n—TR) (Table 6).
Arch7: consisting of two unique regions, U_L and U_S, flanked by the TR_L/IR_L and TR_S/IR_S regions and another region named X located between the IR_L and the IR_S region (i.e. TR_L—U_L—IR_L—X—IR_S—U_S—TR_S) (Table 6).

Table 6.

Representation of the six canonical genomic architectures found in herpesviruses

Name	Genomic structure	Isomers identified	Family
Arch-1	U_L—IR_S—U_S—TR_S	U_S permutation	Alphaherpesvirinae
Arch-2	TR_L—U_L—IR_L—IR_S—U_S—TR_S	U_L and U_S permutations	Alphaherpesvirinae, Betaherpesvirinae
Arch-3	a—TR_L—U_L—IR_L—a—IR_S—U_S—TR_S—a	U_L and U_S permutations	Alphaherpesvirinae
Arch-4	U	-	Betaherpesvirinae, Alloherpesvirinae
Arch-5	TR—U—TR	-	Gammaherpesvirinae, Betaherpesvirinae, Alloherpesvirinae
Arch-6	TR—[U_(n)—IR_(n)]_n—TR	-	Gammaherpesvirinae
Arch-7	TR_L—U_L—IR_L—X—IR_S—U_S—TR_S	U_L and U_S permutations	Malacoherpesvirinae

Open in a new tab

Orthoherpesviridae

The genomic architecture of Orthoherpesviridae viruses varies between families and different genera, as well as within the same genus.

Alphaherpesvirinae

Among the 49 herpesviruses species in the subfamily Alphaherpesvirinae, 45 genomes have been well characterized, two have only partial genomes and two have no nucleotide sequences in public databases (https://ictv.global/report/chapter/orthoherpesviridae/orthoherpesviridae/alphaherpesvirinae, consulted on 28/01/2025). Their genome size ranges from 120 kb for the bird herpesvirus Iltovirus psittacidalpha5, more commonly known as Psittacid alpaherpesvirus 5 (PsHV-5) [84], to 204 kb for the bird herpesvirus Mardivirus columbidalpha1, or Columbid Alphaherpesvirus 1 (CoAHV-1, PHV) [59] (Fig. 3B).

Four different genomic architectures have been described in the subfamily Alphaherpesvirinae: 22 alphaherpesviruses of the genera Scatuvirus, Iltovirus, Mardivirus, Simplexvirus and Varicellovirus share the genomic architecture Arch-1 (Table 6), 11 alphaherpesviruses of the genera Mardivirus, Simplexvirus and Varicellovirus share the genomic architecture Arch-2 (Table 6), 11 alphaherpesviruses of the genera Mardivirus and Simplexvirus share the genomic architecture Arch-3 (Table 6) and the Iltovirus psittacidalpha5 (PsHV-5) [84] have the genomic architecture Arch-4 (Table 6) (Fig. 3B).

Betaherpesvirinae

Among the 27 herpesviruses species of the subfamily Betaherpesvirinae, 24 have a complete genome, while three possess only partial genome. Of the 24 herpesviruses with a complete genome, the genomic arrangement of 19 has been characterized. Their size ranges from 128 kb for the swine herpesvirus Roseolovirus suidbeta2, more commonly known as porcine cytomegalovirus (SuBHV2; PCMV) [164], to 241 kb for the primate herpesvirus Cytomegalovirus paninebeta2, more commonly known as chimpanzee cytomegalovirus (PnBHV2, CCMV) [91] (https://ictv.global/report/chapter/orthoherpesviridae/orthoherpesviridae/betaherpesvirinae, consulted on 28/01/2025).

Three distinct genomic architecture have been described in the subfamily Betaherpesvirinae: three betaherpesviruses of the genus Cytomegalovirus share the genomic architecture Arch-2 (Table 6), one herpesvirus of the genus Quwivirus and one herpesvirus of the genus Muromegalovirus share the genomic architecture Arch-4 (Table 6), and 14 viruses of the genera Cytomegalovirus, Muromegalovirus, Proboscivirus, Quwivirus and Roseolovirus share the genomic architecture Arch-5 (Table 6) (Fig. 3B).

Gammaherpesvirinae

Among the 42 herpesvirus species of the subfamily Gammaherpesvirinae, 26 have a complete genome, whereas 16 possess only a partial genome. Of the 26 herpesviruses species with a complete genome, the genomic architecture of 18 viruses has been characterized. The genome size of these viruses ranges from 119 kb for the Rhadinovirus muridgamma4, better known as Murine gammaherpesvirus 68 (MHV68) [132], to 184 kb for the equine herpesvirus Percavirus equidgamma2, better known as equine herpesvirus 2 (EqGHV2, EVH2) [115] (https://ictv.global/report/chapter/orthoherpesviridae/orthoherpesviridae/gammaherpesvirinae, consulted on 28/01/2025).

Two distinct genomic architectures have been described in Gammaherpesvirinae: 10 viruses of the genera Bossavirus, Macavirus, Percavirus and Rhadinovirus share the genomic architecture Arch-5 (Table 6) whereas 8 gammaherpesviruses of the genus Lymphcryptovirus and Patagavirus share the genomic architecture Arch-6 (Table 6) (Fig. 3B).

Malacoherpesviridae

The complete genomes of the three viruses of the family Malacoherpesviridae have been characterized. With sizes ranging from 192 kb for the Babylonia areolate herpesvirus [19] to 212 kb for the Aurivirus haliotidmalaco 1; better known as Abalone herpesvirus 1 [143] (https://ictv.global/report/chapter/malacoherpesviridae/malacoherpesviridae, consulted on 28/01/2025).

These three malacoherpesviruses share the genomic architecture Arch-7 (Table 6) (Fig. 3B).

Alloherpesviridae

The family Alloherpesviridae contains 13 viruses of which 9 have a complete genome available in genomic public database. Among these 9 herpesviruses, the genomic architecture of 8 viruses have been characterized. The genome sizes of these viruses range from 134 kb for the catfish herpesvirus Ictavirus ictaluridallo1, more commonly known as ictalurid herpesvirus 1 (CCV) [147], to 295 kb for the koi herpesvirus Cyvirus cyprinidallo3, or Cyprinid herpesvirus 3 (CyHV3) [151] (https://ictv.global/report/chapter/alloherpesviridae/alloherpesviridae, consuled on 28/01/2025).

Two distinct genomic architecture have been described in the family Alloherpesviridae: the Batravirus ranidallo3, also known as Ranid herpesvirus 3 (RaHV3) [145] and the bufonid herpesvirus 1 (not classified by ICTV yet) [20] share the genomic architecture Arch-4 (Table 6), and the Lake sturgeon herpesvirus 1 (not classified by ICTV yet) [21] has the genomic architecture Arch-2 (Table 6) whereas all viruses with a complete genome of the genera Cyvirus and Ictavirus, the Silurid herpesvirus 1 (not classified by ICTV yet) [22] and two viruses of the genus Batravirus share the genomic architecture Arch-5 (Table 6) (Fig. 3B).

Insights into the role of herpesvirus genomic architecture

Herpesvirus genomes are among the most complex viral genomes [6]. The genomic architecture of these genomes varies between families and genera. However, the majority of herpesviruses have inverted repeat regions making their genomes difficult to characterise. While the genome architecture of known Malacoherpesviridae viruses has been fully characterized, it is still lacking for some viruses of the Alloherpesviridae, Betaherpesvirinae and Gammaherpesvirinae families. Recent advances in long-read sequencing technologies have provided powerful tools for the sequencing and assembly of complex viral genomes, enabling more accurate and comprehensive analyses of genomic structures. For instance, long-read sequencing has allowed resolving the inverted repeat regions of OsHV-1 [165], HSV-2 [166], and EfHV [135], and detecting genomic isomerization events through sequencing [160, 165–167]. Inverted repeat regions are thought to contribute to the formation of genomic isomers within individual viruses, with potential implications for the regulation of the herpesvirus life cycle [168, 169]. Therefore, a better understanding of these genomic architectures is crucial to characterise and understand their impact on the viral life cycle, viral infectivity and adaptation to their host.

Genomic isomerization of Herpesvirales and its phenotypic implications

The genomic architectures described above represent the canonical genomic organisation of the various herpesviruses characterized to date (Tables 1, 2, 3, 4 and 5). However, several large structural variations termed isomers, resulting from the permutation of the unique regions surrounded by repeated and inverted regions (IR_L/TR_L and IR_S/TR_S) have been described [32, 49, 86, 165–174]. Several approaches have been used to characterise these viral isomerisations, including restriction endonuclease digestion and southern blot hybridisation [32, 49, 86, 167–173], DNA denaturation and electron microscopy [166], long-range PCR [174] and more recently long-read sequencing [160, 165].

Three genomic architectures (Arch-2, Arch-3, and Arch-7) can lead to four viral isomerisation by the permutation of U_L and U_S in the Alphaherpesvirinae, Betaherpesvirinae and Malacoherpesviridae families (Table 6) [32, 47, 49, 50, 50, 51, 54, 86, 165, 172–176]. In addition, the genomic architecture Arch-1 can lead to two viral isomerisations resulting from the permutation of the U_S region, specifically in the Alphaherpesvirinae subfamily [45, 51, 57, 83, 168, 170, 171, 177].

Cause and consequence of genome isomerisation of herpesviruses are still not fully understood. However, for HSV-1, one of the herpesviruses whose isomers have been well studied, isomerisation appears to result from homologous recombination between inverted repeat regions during DNA replication [178, 179]. DNA denaturation and electron microscopy allowed characterizing four HSV-1 genome isomers [166]: (i) the P isomer (for prototype), where U_L and U_S are in sense direction; (ii) the I_L where U_L is inverted; (iii) the I_S where U_S is inverted; and finally (iv) the I_LS, involving inversions of both the U_L and U_S regions. These isomers are typically found in equimolar proportions in HSV-1 [180, 181]. Similar isomer proportion patterns have been observed for HSV-2 [165] and for OsHV-1 [32, 160].

Interestingly, for HSV-1, isomer proportions seem to depend on the type of cells infected and the virus strain. Indeed, in HSV-1 KOS strain isolated from the mouse cornea, the proportions of the four isomers are equivalent. In contrast, when the virus is isolated from the trigeminal ganglia of infected mice, a site where HSV-1 establishes latency, the I_LS isomer is overrepresented [175].

Although, isomers have been identified and characterised for several herpesvirus families, many other herpesviruses including repeated and inverted genomic regions could also show signs of isomerisations. New long read sequencing methods should facilitate the characterisation of herpesvirus isomerisation. Interestingly, analysis of gene annotations across all complete herpesvirus genomes shows that viral genes do not span isomerization boundaries suggesting that gene integrity is preserved during genomic isomerization events. However, the functional consequences of isomerization remain unclear. Further investigation is needed to determine whether different genomic isomers influence viral replication dynamics or infectivity, and whether the coexistence of multiple isomeric forms within the same host cell is required for productive infection. To date, the biological role of genomic isomers remains largely unknown, particularly with respect to their potential impact on replication fidelity and viral fitness.

Herpesviruses life cycle

The viral cycle of herpesviruses is divided into two main phases: a lytic phase and a persistence phase. During the lytic phase, viral genes are expressed and translated into viral proteins that regulate viral DNA replication, the formation of new capsids and the production of new viral particles shed from infected host cells, allowing the virus to replicate and spread to new hosts [182]. During the persistence phase, the virus remains in the host without being damaged by the host's immune system. In general, two scenarios can occur during this phase: the virus can either establish a balance between its replication rate and the host’s immune response, or it can enter a latent state in which viral replication ceases [183–185]. Although all herpesviruses studied for persistence characterisation enter latency, the persistence phase has not been described for all herpesviruses. In the following sections, we will use the term “persistence” (Fig. 3D) to refer to herpesviruses known to establish persistence but for which latency has not been demonstrated or studied at the molecular level, and “latency” (Fig. 3D) to refer to those with demonstrated latency.

Lytic phase

The lytic phase described in this section provides a general overview of the herpesvirus lytic cycle, primarily based on key steps elucidated in extensively studied human herpesviruses. In contrast, the replication cycles of herpesviruses infecting non-human hosts remain largely uncharacterized. To date, among the 137 herpesvirus species, replication has been studied in only 19 of 49 alphaherpesviruses, 9 of 27 betaherpesviruses, 9 of 42 gammaherpesviruses, 2 of 3 malacoherpesviruses, and 3 of 16 alloherpesviruses. This limited knowledge is largely attributable to the difficulty of working with unculturable viruses, as suitable cell lines are unavailable for most host species.

The lytic phase of herpesviruses generally involves several key steps (Fig. 4). First, viral particles bind to receptors on the surface of host cells, a process mediated by glycoproteins located on the surface of the viral envelope (Fig. 4, step 1). Once receptor binding is complete, herpesviruses enter in the cells via two primary mechanisms: direct fusion of the viral envelope with the cell membrane, or membrane fusion following uptake by endocytosis (Fig. 4, step 2) [8, 186, 187]. Once within the host cell, the capsid along with the viral proteins carried by the envelope are released into the cytoplasm. These envelop proteins facilitate the transport of the capsid to the host cell nucleus. In fact, due to the dense and highly organised structure of the cytoplasm, diffusion alone is not sufficient for capsid movement from the cell surface to the nucleus. Instead, capsids actively use microtubules as a transport pathway by binding to microtubule motors through a series of interactions (Fig. 4, step 4). Finally, capsids dock with the nuclear pore complex and release their DNA which is then translocated into the nucleus [8, 25, 188].

Once the viral genome is released into the nucleus, transcription of viral genes begins with the expression of specific viral genes (Fig. 4, step 4). These genes have been classified in three categories depending on the chronology of their expression: (1) immediate-early (IE) or α genes, (2) early (E) or β genes, and (3) late (L) or γ genes [8, 191, 193]. The first genes to be transcribed are the IE genes and code for protein involved in the activation of the transcription of other viral genes [193, 194], RNA splicing [195], and inhibit transcription of host genes [196]. Polypeptides and proteins encoded by E genes are mainly involved in the activation of DNA replication [8, 193], but also in the inhibition of the expression of some IE genes and host cell genes. Finally, late genes are expressed once the genome replication starts [8, 187]. Many of these L genes code for structural proteins involved in virus assembly [8, 25]. At this stage of the infection, the three categories of genes may be expressed simultaneously [25, 26].

DNA replication occurs in the nucleus of infected cells and usually begins when E genes are expressed (Fig. 4, step 5). DNA replication is a two-step process involving initiation by the binding of the origin binding protein to the origin of replication leading to the formation of the replication complex, which replicates the molecule through an intermediate such a theta molecule (Fig. 4, step 5.a) in a process that produces circular progeny DNA molecules. A rolling circle mechanism has been proposed for the second step of DNA synthesis (Fig. 4, step 5b) [8]. This mechanism produces long concatemers of viral genomes (i.e. long DNA molecules consisting of repeated genomes forming a linear multimer), which are cut into genomic units during packaging of the DNA into newly formed capsids [192, 197].

Newly synthesized capsid proteins accumulate in the nucleus to form three types of capsids: C capsids, in which DNA is packaged (Fig. 4, step 6); B capsids, in which viral proteins accumulate; and A capsids which remain empty of DNA or viral proteins and are likely the result of aborted packaging step [25, 26].

The DNA-filled capsids (nucleocapsids) exit the nucleus through a process of envelopment at the inner nuclear membrane forming a primary enveloped particle within the perinuclear space (Fig. 4, step 7.a.). This step is followed by fusion of the primary enveloped particle with the outer nuclear membrane releasing the nucleocapsid without the primary envelop into the cytoplasm with the help of the nuclear egress complex (NEC) (Fig. 4, Step 7.b) [198–201]. The released nucleocapsid then acquires its tegument and mature envelope through different cytoplasmic organelles depending on the herpesvirus species. For example, HCMV tegumentation takes place in the cytoplasmic virion assembly complex (cVAC) [202–204] whereas HSV-1 acquires its tegument from the Golgi apparatus [8]. Tegumented nucleocapsids acquire their mature (secondary) envelope either in the trans-Golgi network for HSV-1, recycling endosomes for HCMV or multivesicular bodies for HHV-6A (Fig. 4, step 7.c) [8, 198, 199]. Vesicles containing enveloped virions are transported to the plasma membrane, where vesicle membrane fuses with the plasma membrane. Mature virions are then released into the extracellular space where they can infect other cells from the same or from different individuals (Fig. 4, step 8) [8, 25, 198, 199].

Persistent phase

The second phase of the herpesvirus life cycle is the persistence phase. Among the 137 recognized herpesvirus species, persistence has been investigated in 28 of 49 alphaherpesviruses, 10 of 27 betaherpesviruses, 17 of 42 gammaherpesviruses, 1 of 3 malacoherpesviruses, and 6 of 13 alloherpesviruses. However, molecular evidence of latency has been demonstrated for only 31 of these 62 species (Fig. 3D) [49, 133, 140, 141, 205–234].

Herpesviruses that have been characterized for persistence primarily persist in two specific tissues within their hosts. Herpesviruses of the family Alphaherpesvirinae establish persistence primarily in the trigeminal ganglion of their hosts (23 herpesviruses species, Fig. 3D) [47, 67, 80, 211, 214, 223, 230, 235–245] (Fig. 3D). However, Galliformes herpesviruses and herpesviruses of the subfamilies Betaherpesvirinae, Gammaherpesvirinae, and of the families Malacoherpesviridae and Alloherpesviridae establish their persistence in the immune system cells of their hosts (28 herpesviruses species, Fig. 3D) [24, 140, 216, 224, 225, 228, 246–248] (in green in Fig. 3D).

Based on a comprehensive literature review of 31 herpesviruses with documented latency at the molecular level, the following sections describe the mechanisms that maintain viral genome and gene expression during latency and the processes that trigger herpesvirus reactivation.

Maintenance of the viral genome during latency

Depending on the host, different mechanisms are required to maintain latency. For herpesviruses that establish latency in dividing cells, these mechanisms must ensure stable retention of the viral genome in the nucleus of infected cells and its transmission to daughter cells during cell division. There is relatively little information in the literature about these mechanisms. To date, two main mechanisms have been described to maintain viral herpes genome in infected cells: the circularisation of the viral genome as episome (10 herpesviruses species, Fig. 3D) or the integration of the viral genome into host DNA (4 herpesviruses species, Fig. 3D).

Episomes formation has been demonstrated for several herpesviruses including human herpesviruses (i.e. HSV-1, HSV-2, Cytomegalovirus humanbeta5 (HCMV), Varicellovirus humanalpha3 (VZV), Lymphocryptovirus humangamma4 (EBV), HHV-6, and Roseolovirus humanbeta7 HHV-7) [24], bovines herpesvirus (Macavirus ovinegamma2, OvGHV-2) [228, 249], bat herpesvirus (Patagivirus vespertilionidgamma3, EfHV) [250], mouse herpesvirus (Rhadinovirus muridgamma4, MHV68) [251] and fish herpesvirus (i.e. Ictavirus ictaluridallo1, IcHV1) [227] (Fig. 3D). After initial infection and virus entry into the trigeminal ganglion or host immune cells, the linear herpesvirus genome becomes circularised by fusion of terminal repeats located at both ends of the genome and is rapidly chromatinised in the nucleus [24, 252–254, 254, 255]. Herpesviruses that establish latency in immune cells capable of division must ensure their persistence during cell replication (Fig. 3D). To achieve this, they synthesize viral proteins that facilitate the attachment of the viral episome to the host chromosome, enabling the transmission of the viral genome to daughter cells. [256] (Fig. 5A).

Integration into host DNA has been described for MDV, HHV-6 and EBV (Fig. 5B). The process of integration involves the terminal regions of MDV and HHV-6 genomes that contain perfect repeats of the telomeric sequence of their host chromosomes [257–259]. While the precise mechanisms underlying this integration remain unclear, certain viral proteins such as 5’-3’ exonuclease, single-strand DNA binding protein or adeno-associated virus 2 integrase, may facilitate MDV and HHV-6 integration, respectively [260]. In contrast, EBV does not integrate at a single site like MDV and HHV-6; instead, its integration is non-random, occurring predominantly in heterochromatic regions of the host genome, which are rich in repetitive sequences and lack functional genes [261]. By integrating into the host DNA, these herpesviruses ensure the stable maintenance of their genome during cell division.

Regulation of viral genes expression during latency

To evade detection by the host immune system, herpesviruses must tightly regulate the expression of their lytic genes and modulate host genes activity to prevent elimination of the infected cell. This is achieved by three different mechanisms including expression of viral long non-coding RNAs (lncRNA), microRNAs (miRNA) and non-lytic protein-coding genes.

Many herpesviruses express lncRNAs that contribute to the establishment of latency. These transcripts are transcribed antisense to immediate-early genes and induce the regulation of their transcription [24, 231, 262, 263]. lncRNAs are also involved in epigenetic modifications of histones, such as trimethylation and demethylation, leading to the formation of repressive heterochromatin around viral DNA, thereby preventing lytic gene transcription. In addition, these lncRNAs contribute to the inhibition of apoptosis, ensuring the long-term persistence of the viral genome (Fig. 6) [24, 262].

Fig. 6 — Mechanisms of regulation of viral genes expression during latency.LncRNAs, protein-coding genes and miRNAs are expressed during the latency phase and play a role in the maintenance of the latency by modulating histone epigenetics, inhibiting apoptosis of the infected cells, stimulating the cell cycle progression, blocking translation of lytic proteins, regulating viral and host expression and suppressing lytic gene expression

Viral and host miRNAs play an important role in modulating latency [24, 264, 265]. They can contribute to the degradation of lytic mRNA or to the inhibition of lytic protein translation, and to the regulation of host cell genes expression. All these mechanisms lead to the evasion of recognition of latently infected cells by the host immune system, to reduce the expression of immediate-early genes and are also involved in the inhibition of apoptosis [24, 264, 266, 267] (Fig. 6).

Finally, some herpesviruses express protein-coding genes that play also essential roles in maintaining latency. Similar to lncRNAs and miRNAs, these non-lytic proteins are involved in epigenetic modifications of histone, tethering the viral episome to the host chromosome, promoting cell cycle progression, and inhibiting apoptosis. Together, all these processes contribute to the stable persistence of the viral genome inside host cells (Fig. 6) [24, 209, 268].

While these mechanisms ensure the long-term viral latency, herpesviruses have also developed strategies to emerge from this dormant state upon stress activation and when conditions are favourable for viral replication.

Reactivation

The success of herpesviruses is primarily due to their ability to reactivate from a latent state. Viral reactivation can occur spontaneously, but is more likely to occur in response to biotic or abiotic stress of their host. In the case of HSV-1, several stimuli that induce neuronal stress have been shown to trigger viral reactivation in cell culture, including removal or inhibition of neurotrophic factors and exposure to UV light. In humans, triggers include exposure to sunlight, hormonal fluctuations or surgical resection [193, 269–273]. For Varicellovirus felidalpha1 (FVRV) which infects cats, stressors such as environmental changes can lead to viral reactivation [220], while for MDV: Mardivirus gallidalpha2, also known as Marek’s disease virus (MDV), factors such as hypoxia, apoptosis or a decrease of temperature can induce viral reactivation in cell culture [24, 251, 274] (Fig. 5A). For herpesviruses that integrate inside telomeres of their host chromosome, viral reactivation could also be induced by the natural shortening of the telomeres that could either relieve suppression of viral gene transcription or inducing telomere-telomere recombination, resulting in release of the virus genome from the integration site (Fig. 5B) [260].

The mechanisms of herpesviruses reactivation are not fully understood. Based on the work done on HSV-1 and murine neurons, the reactivation can be divided in two phases. The first one, called “animation”, involves decrease of lncRNAs and miRNAs expression, transcription from the silenced DNA that does not follow the same well-ordered process of gene transcription that occurs during the lytic phase [193, 253, 263, 275, 276]. Once some viral proteins are translated, the second reactivation phase starts, classical cascade of virus gene expression occurs (see Lytic Phase), progeny virions are assembled and transported to axon termini to be released in epithelial cells in the mucosa or skin resulting in disease [24, 193, 276].

Perspectives and insights into herpesvirus life cycle comprehension

For most herpesvirus species (95 out of 137), the lytic phase has not yet been described at the transcriptomic level. This is likely due to the high transcriptional complexity of herpesviruses [277], as well as challenges in developing suitable models to study them and/or difficulties in inducing the lytic phase in some species [278]. Latency has been demonstrated in 31 of the 137 known herpesvirus species but it is likely that other herpesviruses also have the ability to establish latency in their hosts. With the exception of the Malacoherpesviridae family, all herpesvirus families contain at least one species for which latency has been demonstrated at the molecular level, supporting the hypothesis that latency is a common feature of the persistent phase. Indeed, aquatic herpesviruses become undetectable at low water temperatures in winter and reactivate in summer when temperatures rise, suggesting the presence of a latency phase [279].

Recent advances in sequencing and imagery technologies, such as single-cell genomics and transcriptomics, spatial transcriptomic and multi-omics approaches, allow studying viral gene expression alongside the host cell immune response, providing a more comprehensive view of the viral development in different tissues or even in different cell types [280, 281]. Long-read sequencing technologies enable the characterization of full-length viral transcripts and the identification of polycistronic RNAs [282, 283]. Moreover, the study of epigenetic modifications [284–287] could also provide new insights into the life cycle of herpesvirus. This may not only help to characterize the production of new virions at the transcriptomic level, but also to identify non-coding viral transcripts that may support latency.

In addition, many viral genes have not yet been fully functionally annotated, and their true coding potential is still poorly understood. Transcriptomics coupled with proteomics analyses could overcome these limits [283, 288]. Furthermore, machine learning tools such as AlphaFold [289] and DeepLoc (https://services.healthtech.dtu.dk/services/DeepLoc-2.1/) could be used to investigate virus-host protein interactions as well as to predict the localization of the protein in the host cell.

These recent advances in sequencing technologies and bioinformatics tools should significantly improve our knowledge of herpesvirus biology in the coming years.

Conclusion

This review offers an overview on the available information on systematics, genomic architecture and life cycle of herpesviruses. Herpesviruses are large and complex double-stranded DNA viruses, characterized by a capsid that encloses tightly packed DNA, surrounded by a protein-rich tegument and an outer envelope. They have a complex, family-specific genomic architecture that can be divided into seven groups. For almost all herpesviruses, the genomic architecture includes inverted repeat regions, that lead to the formation of isomers that coexist in equimolar proportions within host cells. The function of these isomers remains poorly understood, probably due to the challenges of their characterization.

To date, 137 herpesviruses species have been described, but many more, particularly within the Malacoherpesviridae and Alloherpesviridae families, are likely to remain undiscovered. Advances in database analysis, long-read sequencing and purification methods may help facilitate their characterization.

The life cycle of human-infecting herpesviruses is well documented, showing a sequential expression of three categories of genes required for the assembly of new viral particles. In contrast, the replication cycles of non-human herpesviruses remain poorly understood largely due to the lack of suitable cell lines for many non-human species, making in vitro studies difficult. Similar limitations apply to the study of latency at the molecular level. Although several long non-coding RNAs (lncRNAs) and microRNAs (miRNAs) involved in maintaining the viral genome during latency, either integrated or circularized as an episome, have been identified in human herpesviruses, much less is known about these processes in other hosts. This knowledge gap is particularly evident for members of the families Gammaherpesviridae, Malacoherpesviridae, and Alloherpesviridae.

Improving our understanding of the genomic architecture, isomeric configurations and life cycles of herpesviruses, especially those that infect non-human hosts, will be crucial to fill current knowledge gaps. Recent breakthroughs in genomics, transcriptomics, proteomics and imaging now provide the tools to explore these aspects in greater depth and unravel the complex biology and evolutionary dynamics of these viruses, ultimately helping to develop strategies to prevent reactivation and associated disease.

Acknowledgements

Not applicable.

Abbreviations

AbHV-1: Aurivirus haliotidmalaco1
BaHV: Babylonia areolate Herpesvirus
CCV: Ictavirus ictaluridallo1
CoAHV-1, PHV: Mardivirus columbidalpha1
cVAC: Cytoplasmic Virion Assembly complex
CyHV3: Cyvirus cyprinidallo3
dsDNA: Double-stranded DNA
dsRNA: Double-stranded RNA
E: Early
EBV: Lymphocryptovirus humangamma4
EfHV: Patagivirus vespertilionidgamma3
EqGHV2, EHV2: Percavirus equidgamma2
FRV: Varicellovirus felidalpha1
HCMV: Cytomegalovirus humanbeta5
HHV-6A: Roseolovirus humanbeta6
HHV-7: Roseolovirus humanbeta7
HSV-1: Simplexvirus humanalpha1
HSV-2: Simplexvirus humanalpha2
IcHV1: Ictavirus ictaluridallo1
ICTV: International Committee on Taxonomy of Viruses
IE: Immediate-early
IR: Internal Repeat
IR_L: Internal Repeat Long
IR_S: Internal Repeat Short
L: Late
lncRNA: Long non-coding RNA
MDV: Mardivirus gallidalpha2
MHV68: Rhadinovirus muridgamma4
miRNA: Micro RNA
NEC: Nuclear Egress Complex
NGS: Next Generation Sequencing
OcGHV2: Macavirus ovinegamma2
OsHV-1: Ostreavirus ostreidmalaco1
PnBHV2, CCMV: Cytomegalovirus paninebeta2
PsHV-5: Iltovirus psittacidalpha5
RaHV3: Batravirus ranidallo3
ssDNA: Single-stranded DNA
ssRNA: Single-stranded RNA
SuBHV2, PMCV: Roseolovirus suidbeta2
TR: Terminal Repeat
TR_L: Terminal Repeat Long
TR_S: Terminal Repeat Short
U_L: Unique Long
U_S: Unique Short
VZV: Varicellovirus humanalpha3

Authors’ contributions

ADM drafted the manuscript and GC, BM and IA corrected it. All authors read and approved the final version of the manuscript.

Funding

ADM, GC, BM and IA were financially supported by Ifremer.

Data availability

No datasets were generated or analysed during the current study.

Declarations

Ethics approval and consent to participate

Not applicable.

Consent for publication

Not applicable.

Competing interests

The authors declare no competing interests.

Footnotes

Publisher’s Note

Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Contributor Information

Aurélie Dotto-Maurel, Email: aurelie.dotto.maurel@ifremer.fr.

Germain Chevignon, Email: germain.chevignon@ifremer.fr.

References

1.Edwards RA, Rohwer F. Viral metagenomics. Nat Rev Microbiol juin. 2005;3(6):504–10. [DOI] [PubMed] [Google Scholar]
2.Hendrix RW, Smith MC, Burns RN, Ford ME, Hatfull GF. Evolutionary relationships among diverse bacteriophages and prophages: all the world’s a phage. Proc Natl Acad Sci U S A. 1999;96(5):2192–7. [DOI] [PMC free article] [PubMed] [Google Scholar]
3.Lawrence CM, Menon S, Eilers BJ, Bothner B, Khayat R, Douglas T, et al. Structural and functional studies of archaeal viruses. J Biol Chem. 2009;284(19):12599–603. [DOI] [PMC free article] [PubMed] [Google Scholar]
4.Nayfach S, Camargo AP, Schulz F, Eloe-Fadrosh E, Roux S, Kyrpides NC. CheckV assesses the quality and completeness of metagenome-assembled viral genomes | Nature Biotechnology. 2021. Disponible sur: https://www.nature.com/articles/s41587-020-00774-7. Cité 17 Nov 2024. [DOI] [PMC free article] [PubMed]
5.O’Leary NA, Wright MW, Brister JR, Ciufo S, Haddad D, McVeigh R, et al. Reference sequence (RefSeq) database at NCBI: current status, taxonomic expansion, and functional annotation. Nucleic Acids Res. 2016;44(D1):D733-745. [DOI] [PMC free article] [PubMed] [Google Scholar]
6.Silva JM, Pratas D, Caetano T, Matos S. The complexity landscape of viral genomes. GigaScience. 2022;11:giac079. [DOI] [PMC free article] [PubMed] [Google Scholar]
7.Gatherer D, Depledge DP, Hartley CA, Szpara ML, Vaz PK, Benkő M, et al. ICTV virus taxonomy profile: herpesviridae 2021. J Gen Virol. 2021;102(10):001673. [DOI] [PMC free article] [PubMed] [Google Scholar]
8.Knipe DM, Howley PM, Cohen JI, Griffin DE, Lamb RA, Martin MA, et al. Fields virology 6th edition. 2013.
9.Brito AF, Baele G, Nahata KD, Grubaugh ND, Pinney JW. Intrahost speciations and host switches played an important role in the evolution of herpesviruses. Virus Evol. 2021;7(1):veab025. [DOI] [PMC free article] [PubMed] [Google Scholar]
10.McGeoch DJ, Cook S, Dolan A, Jamieson FE, Telford EA. Molecular phylogeny and evolutionary timescale for the family of mammalian herpesviruses. J Mol Biol. 1995;247(3):443–58. [DOI] [PubMed] [Google Scholar]
11.McGeoch DJ, Dolan A, Ralph AC. Toward a comprehensive phylogeny for mammalian and avian herpesviruses. J Virol. 2000;74(22):10401–6. [DOI] [PMC free article] [PubMed] [Google Scholar]
12.McGeoch DJ, Cook S. Molecular phylogeny of the alphaherpesvirinae subfamily and a proposed evolutionary timescale. J Mol Biol. 1994;238(1):9–22. [DOI] [PubMed] [Google Scholar]
13.Davison AJ. Evolution of the herpesviruses. Vet Microbiol. 2002;86(1–2):69–88. [DOI] [PubMed] [Google Scholar]
14.Jackson A. The effect of paralogous lineages on the application of reconciliation analysis by cophylogeny mapping. Syst Biol. 2005;54(1):127–45. [DOI] [PubMed] [Google Scholar]
15.McGeoch DJ, Rixon FJ, Davison AJ. Topics in herpesvirus genomics and evolution. Virus Res. 2006;117(1):90–104. [DOI] [PubMed] [Google Scholar]
16.Escalera-Zamudio M, Rojas-Anaya E, Kolokotronis SO, Taboada B, Loza-Rubio E, Méndez-Ojeda ML, et al. Bats, primates, and the evolutionary origins and diversification of mammalian gammaherpesviruses. mBio. 2016;7(6):e01425. 10.1128/mbio.01425-16. [DOI] [PMC free article] [PubMed] [Google Scholar]
17.McGeoch DJ, Davison AJ, Dolan A, Gatherer D, Sevilla-Reyes EE. Molecular Evolution of the Herpesvirales. In: Domingo E, Parrish CR, Holland JJ, éditeurs. Origin and Evolution of Viruses (Second Edition). London: Academic Press; 2008. p. 447‑75. Disponible sur: https://www.sciencedirect.com/science/article/pii/B9780123741530000205. Cité 22 Janv 2025.
18.Fu M, Deng R, Wang J, Wang X. Whole-genome phylogenetic analysis of herpesviruses. Acta Virol. 2008;52(1):31–40. [PubMed] [Google Scholar]
19.Divilov K. Whole-genome assembly of a novel invertebrate herpesvirus from the gastropod Babylonia areolata. Microb Genom. 2024;10(4):001237. [DOI] [PMC free article] [PubMed] [Google Scholar]
20.Origgi FC, Schmidt BR, Lohmann P, Otten P, Meier RK, Pisano SRR, et al. Bufonid herpesvirus 1 (BfHV1) associated dermatitis and mortality in free ranging common toads (Bufo bufo) in Switzerland. Sci Rep. 2018;8(1):14737. [DOI] [PMC free article] [PubMed] [Google Scholar]
21.Walker L, Subramaniam K, Viadanna PHO, Vann JA, Marcquenski S, Godard D, et al. Characterization of an alloherpesvirus from wild lake sturgeon Acipenser fulvescens in Wisconsin (USA). Dis Aquat Organ. 2022;149:83–96. [DOI] [PubMed] [Google Scholar]
22.Henriquez RN, Polchana J, Kanchanakhan S, Davison AJ, Waltzek TB, Subramaniam K. Genome sequence of a novel alloherpesvirus isolated from glass catfish (Kryptopterus bicirrhis). Genome Announc. 2018;6(19):e00403. 10.1128/genomea.00403-18. [DOI] [PMC free article] [PubMed] [Google Scholar]
23.Davison AJ, Clements JB. Herpesviruses: General Properties. In: Topley & Wilson’s Microbiology and Microbial Infections. John Wiley & Sons, Ltd; 2010. Disponible sur: https://onlinelibrary.wiley.com/doi/abs/10.1002/9780470688618.taw0231. Cité 15 Jan 2025.
24.Cohen JI. Herpesvirus latency. American Society for Clinical Investigation; 2020. Disponible sur: https://www.jci.org/articles/view/136225/pdf. Cité 5 Jan 2023.
25.Everett RD. HSV-1 Biology and Life Cycle. In: Diefenbach RJ, Fraefel C, éditeurs. Herpes Simplex Virus: Methods and Protocols. New York, NY: Springer; 2014. p. 1‑17. (Methods in Molecular Biology). Disponible sur: 10.1007/978-1-4939-0428-0_1. Cité 17 juill 2023.
26.Griffin DE, Knipe D, Howley P, Lamb R, Martin M, Roizman B, et al. Fields virology. 2001.
27.Davison AJ. Herpesvirus systematics. Vet Microbiol. 2010;143(1):52–69. [DOI] [PMC free article] [PubMed] [Google Scholar]
28.Pellett PE, Roizman B. Chapter 59 : Herpesviridae. In: Fields Virology 6th edition. 2013.
29.Epstein MA. Observations on the mode of release of herpes virus from infected HeLa cells. J Cell Biol. 1962;12(3):589–97. [DOI] [PMC free article] [PubMed] [Google Scholar]
30.McCombs RM, Brunschwig P, Mirkovic R, Benyesh-Melnick M. Electron microscopic characterization of a herpeslike virus isolated from tree shrews. Virology. 1971;45(3):816–20. 10.1016/0042-6822(71)90203-0. [DOI] [PubMed]
31.Booy FP, Trus BL, Davison AJ, Steven AC. The capsid architecture of channel catfish virus, an evolutionarily distant herpesvirus, is largely conserved in the absence of discernible sequence homology with herpes simplex virus. Virology. 1996;215(2):134–41. [DOI] [PubMed] [Google Scholar]
32.Davison AJ, Trus BL, Cheng N, Steven AC, Watson MS, Cunningham C, et al. A novel class of herpesvirus with bivalve hosts. J Gen Virol. 2005;86(1):41–53. [DOI] [PubMed] [Google Scholar]
33.Liu F, Zhou ZH. Comparative virion structures of human herpesviruses. In: Arvin A, Campadelli-Fiume G, Mocarski E, Moore PS, Roizman B, Whitley R, et al., éditeurs. Human Herpesviruses: Biology, Therapy, and Immunoprophylaxis. Cambridge: Cambridge University Press; 2007. Disponible sur: http://www.ncbi.nlm.nih.gov/books/NBK47399/. Cité 18 Nov 2024.
34.Furlong D, Swift H, Roizman B. Arrangement of herpesvirus deoxyribonucleic acid in the core. J Virol. 1972;10(5):1071–4. [DOI] [PMC free article] [PubMed] [Google Scholar]
35.Nazerian K. DNA configuration in the core of Marek’s disease virus. J Virol mai. 1974;13(5):1148–50. [DOI] [PMC free article] [PubMed] [Google Scholar]
36.Zhou ZH, Chen DH, Jakana J, Rixon FJ, Chiu W. Visualization of tegument-capsid interactions and DNA in intact herpes simplex virus type 1 virions. J Virol. 1999;73(4):3210–8. [DOI] [PMC free article] [PubMed] [Google Scholar]
37.Heming JD, Conway JF, Homa FL. Herpesvirus Capsid Assembly and DNA Packaging. In: Osterrieder K, éditeur. Cell Biology of Herpes Viruses. Cham: Springer International Publishing; 2017. p. 119‑42. (Advances in Anatomy, Embryology and Cell Biology; vol. 223). Disponible sur: http://link.springer.com/10.1007/978-3-319-53168-7_6. Cité 20 avr 2023. [DOI] [PMC free article] [PubMed]
38.Johannsen E, Luftig M, Chase MR, Weicksel S, Cahir-McFarland E, Illanes D, et al. Proteins of purified Epstein-Barr virus. Proc Natl Acad Sci U S A. 2004;101(46):16286–91. [DOI] [PMC free article] [PubMed] [Google Scholar]
39.Kattenhorn LM, Mills R, Wagner M, Lomsadze A, Makeev V, Borodovsky M, et al. Identification of proteins associated with murine cytomegalovirus virions. J Virol. 2004;78(20):11187–97. [DOI] [PMC free article] [PubMed] [Google Scholar]
40.Michael K, Klupp BG, Mettenleiter TC, Karger A. Composition of pseudorabies virus particles lacking tegument protein US3, UL47, or UL49 or envelope glycoprotein E. J Virol. 2006;80(3):1332–9. [DOI] [PMC free article] [PubMed] [Google Scholar]
41.O’Connor CM, Kedes DH. Mass spectrometric analyses of purified rhesus monkey rhadinovirus reveal 33 virion-associated proteins. J Virol. 2006;80(3):1574–83. [DOI] [PMC free article] [PubMed] [Google Scholar]
42.Varnum SM, Streblow DN, Monroe ME, Smith P, Auberry KJ, Paša-Tolić L, et al. Identification of Proteins in Human Cytomegalovirus (HCMV) particles: the HCMV proteome. J Virol. 2004;78(20):10960–6. [DOI] [PMC free article] [PubMed] [Google Scholar]
43.Yuan S, Wang J, Zhu D, Wang N, Gao Q, Chen W, et al. Cryo-EM structure of a herpesvirus capsid at 3.1 Å. Science. 2018;360(6384):eaao7283. [DOI] [PubMed] [Google Scholar]
44.Wörner TP, Shamorkina TM, Snijder J, Heck AJR. Mass spectrometry-based structural virology. Anal Chem. 2021;93(1):620–40. [DOI] [PMC free article] [PubMed] [Google Scholar]
45.Eberle R, Black DH. Sequence of the ateline alphaherpesvirus 1 (HVA1) genome. Arch Virol. 2017;162(5):1423–5. [DOI] [PMC free article] [PubMed] [Google Scholar]
46.Pfaff F, Neubauer-Juric A, Krebs S, Hauser A, Singer S, Blum H, et al. Full genome sequence of bovine alphaherpesvirus 2 (BoHV-2). Arch Virol. 2021;166(2):639–43. [DOI] [PMC free article] [PubMed] [Google Scholar]
47.Tyler SD, Peters GA, Severini A. Complete genome sequence of cercopithecine herpesvirus 2 (SA8) and comparison with other simplexviruses. Virology. 2005;331:429. [DOI] [PubMed] [Google Scholar]
48.Davison AJ. Evolution of sexually transmitted and sexually transmissible human herpesviruses. Ann N Y Acad Sci. 2011;1230(1):E37-49. [DOI] [PubMed] [Google Scholar]
49.Babra B, Watson G, Xu W, Jeffrey BM, Xu JR, Rockey DD, et al. Analysis of the genome of leporid herpesvirus 4. Virology. 2012;433(1):183–91. [DOI] [PMC free article] [PubMed] [Google Scholar]
50.Perelygina L, Zhu L, Zurkuhlen H, Mills R, Borodovsky M, Hilliard JK. Complete sequence and comparative analysis of the genome of herpes B virus (Cercopithecine Herpesvirus 1) from a Rhesus Monkey. J Virol. 2003;77(11):6167–77. [DOI] [PMC free article] [PubMed] [Google Scholar]
51.Vaz PK, Mahony TJ, Hartley CA, Fowler EV, Ficorilli N, Lee SW, et al. The first genome sequence of a metatherian herpesvirus: macropodid herpesvirus 1. BMC Genomics. 2016;17(1):70. [DOI] [PMC free article] [PubMed] [Google Scholar]
52.Vaz PK, Mahony TJ, Hartley CA, Motha J, Devlin JM. Genome Sequences of Two Marsupial Simplex Viruses, Macropodid Alphaherpesviruses 2 and 4. Microbiol Resour Announc. 2021;10(2):e01189-20 Matthijnssens J, éditeur. [DOI] [PMC free article] [PubMed] [Google Scholar]
53.Severini A, Tyler SD, Peters GA, Black D, Eberle R. Genome sequence of a chimpanzee herpesvirus and its relation to other primate alphaherpesviruses. Arch Virol. 2013;158(8):1825–8. [DOI] [PMC free article] [PubMed] [Google Scholar]
54.Tyler SD, Severini A. The complete genome sequence of Herpesvirus Papio 2 (Cercopithecine Herpesvirus 16) shows evidence of recombination events among various progenitor herpesviruses. J Virol. 2006;80(3):1214–21. [DOI] [PMC free article] [PubMed] [Google Scholar]
55.Sasaki M, Setiyono A, Handharyani E, Kobayashi S, Rahmadani I, Taha S, et al. Isolation and characterization of a novel alphaherpesvirus in fruit bats. J Virol. 2014;88(17):9819–29. [DOI] [PMC free article] [PubMed] [Google Scholar]
56.Inagaki T, Yamada S, Fujii H, Yoshikawa T, Shibamura M, Harada S, et al. Characterization of a novel alphaherpesvirus isolated from the fruit bat pteropus lylei in Vietnam. J Virol. 2020;94(18):e00673-e720. [DOI] [PMC free article] [PubMed] [Google Scholar]
57.Tyler S, Severini A, Black D, Walker M, Eberle R. Structure and sequence of the saimiriine herpesvirus 1 genome. Virology. 2011;410(1):181–91. [DOI] [PMC free article] [PubMed] [Google Scholar]
58.Wang J, Höper D, Beer M, Osterrieder N. Complete genome sequence of virulent duck enteritis virus (DEV) strain 2085 and comparison with genome sequences of virulent and attenuated DEV strains. Virus Res. 2011;160(1):316–25. [DOI] [PubMed] [Google Scholar]
59.Guo Y, Li S, Sun X, He Y, Zhao H, Wang Y, et al. Complete genome sequence and evolution analysis of a columbid herpesvirus type 1 from feral pigeon in China. Arch Virol. 2017;162(7):2131–3. [DOI] [PubMed] [Google Scholar]
60.Tulman ER, Afonso CL, Lu Z, Zsak L, Rock DL, Kutish GF. The genome of a very virulent Marek’s disease virus. J Virol. 2000;74(17):7980–8. [DOI] [PMC free article] [PubMed] [Google Scholar]
61.Spatz SJ, Schat KA. Comparative genomic sequence analysis of the Marek’s disease vaccine strain SB-1. Virus Genes. 2011;42(3):331–8. [DOI] [PubMed] [Google Scholar]
62.Afonso CL, Tulman ER, Lu Z, Zsak L, Rock DL, Kutish GF. The genome of turkey herpesvirus. J Virol. 2001;75(2):971–8. [DOI] [PMC free article] [PubMed] [Google Scholar]
63.Pfaff F, Schulze C, König P, Franzke K, Bock S, Hlinak A, et al. A novel alphaherpesvirus associated with fatal diseases in banded Penguins. J Gen Virol. 2017;98(1):89–95. [DOI] [PubMed] [Google Scholar]
64.d’Offay JM, Fulton RW, Eberle R. Complete genome sequence of the NVSL BoHV-1.1 Cooper reference strain. Arch Virol. 2013;158(5):1109–13. [DOI] [PubMed] [Google Scholar]
65.Delhon G, Moraes MP, Lu Z, Afonso CL, Flores EF, Weiblen R, et al. Genome of bovine herpesvirus 5. J Virol. 2003;77(19):10339–47. [DOI] [PMC free article] [PubMed] [Google Scholar]
66.Scheffer CM, Varela APM, Cibulski SP, Schmidt C, Campos FS, Paim WP, et al. Genome sequence of bubaline alphaherpesvirus 1 (BuHV1) isolated in Australia in 1972. Arch Virol. 2017;162(5):1169–76. [DOI] [PubMed] [Google Scholar]
67.Papageorgiou KV, Suárez NM, Wilkie GS, McDonald M, Graham EM, Davison AJ. Genome sequence of canine herpesvirus. PLoS One. 2016;11(5):e0156015. [DOI] [PMC free article] [PubMed] [Google Scholar]
68.Hao F, Mao L, Li W, Li J, Yang L, Zhang W, et al. Epidemiological investigation and genomic characterization of Caprine herpesvirus 1 from goats in China. Infect Genet Evol. 2020;79:104168. [DOI] [PubMed] [Google Scholar]
69.Gray WL, Starnes B, White MW, Mahalingam R. The DNA sequence of the simian varicella virus genome. Virology. 2001;284(1):123–30. [DOI] [PubMed] [Google Scholar]
70.Telford EAR, Watson MS, McBride K, Davison AJ. The DNA sequence of equine herpesvirus-1. Virology. 1992;189(1):304–16. [DOI] [PubMed] [Google Scholar]
71.Sijmons S, Vissani A, Tordoya MS, Muylkens B, Thiry E, Maes P, et al. Complete genome sequence of equid Herpesvirus 3. Genome Announc. 2014;2(5):e00797-e814. [DOI] [PMC free article] [PubMed] [Google Scholar]
72.Telford EA, Watson MS, Perry J, Cullinane AA, Davison AJ. The DNA sequence of equine herpesvirus-4. J Gen Virol. 1998;79(5):1197–203. [DOI] [PubMed] [Google Scholar]
73.Martella V, Lanave G, Camero M, Larocca V, Lorusso E, Catella C, et al. Identification of a novel α-herpesvirus associated with ulcerative stomatitis in donkeys. Emerg Infect Dis. 2020;26(12):3044–7. [DOI] [PMC free article] [PubMed] [Google Scholar]
74.Garvey M, Suárez NM, Kerr K, Hector R, Moloney-Quinn L, Arkins S, et al. Equid herpesvirus 8: Complete genome sequence and association with abortion in mares. PLoS One. 2018;13(2):e0192301. [DOI] [PMC free article] [PubMed] [Google Scholar]
75.Kasem S, Yamada S, Kiupel M, Woodruff M, Ohya K, Fukushi H. Equine herpesvirus type 9 in giraffe with encephalitis. Emerg Infect Dis. 2008;14(12):1948–9. [DOI] [PMC free article] [PubMed] [Google Scholar]
76.Tai SHS, Niikura M, Cheng HH, Kruger JM, Wise AG, Maes RK. Complete genomic sequence and an infectious BAC clone of feline herpesvirus-1 (FHV-1). Virology. 2010;401(2):215–27. [DOI] [PubMed] [Google Scholar]
77.Davison AJ, Scott JE. The complete DNA sequence of varicella-zoster virus. J Gen Virol. 1986;67(Pt 9):1759–816. [DOI] [PubMed] [Google Scholar]
78.Davison AJ, Nielsen O, Subramaniam K, Jacob JM, Romero CH, Burek-Huntington KA, et al. Genome Sequence of an Alphaherpesvirus from a Beluga Whale (Delphinapterus leucas). Genome Announc. 2017;5(42):e01100-17. 10.1128/genomea.01100-17. [DOI] [PMC free article] [PubMed] [Google Scholar]
79.van Boheemen S, van Rijn AL, Pappas N, Carbo EC, Vorderman RHP, Sidorov I, et al. Retrospective validation of a metagenomic sequencing protocol for combined detection of RNA and DNA viruses using respiratory samples from pediatric patients. J Mol Diagn. 2020;22(2):196–207. [DOI] [PMC free article] [PubMed] [Google Scholar]
80.Szpara ML, Tafuri YR, Parsons L, Shamim SR, Verstrepen KJ, Legendre M, et al. A wide extent of inter-strain diversity in virulent and vaccine strains of alphaherpesviruses. PLoS Pathog. 2011;7(10):e1002282 Baines JD, éditeur. [DOI] [PMC free article] [PubMed] [Google Scholar]
81.Sutherland M, Sarker S, Vaz PK, Legione AR, Devlin JM, Macwhirter PL, et al. Disease surveillance in wild Victorian cacatuids reveals co-infection with multiple agents and detection of novel avian viruses. Vet Microbiol. 2019;235:257–64. [DOI] [PubMed] [Google Scholar]
82.Lee SW, Devlin JM, Markham JF, Noormohammadi AH, Browning GF, Ficorilli NP, et al. Comparative analysis of the complete genome sequences of two Australian origin live attenuated vaccines of infectious laryngotracheitis virus. Vaccine. 2011;29(52):9583–7. [DOI] [PubMed] [Google Scholar]
83.Thureen D, Keeler C. Psittacid Herpesvirus 1 and infectious laryngotracheitis virus: comparative genome sequence analysis of two avian alphaherpesviruses. J Virol. 2006;80:7863–72. [DOI] [PMC free article] [PubMed] [Google Scholar]
84.Sutherland M, Sarker S, Raidal SR. Molecular and microscopic characterisation of a novel pathogenic herpesvirus from Indian ringneck parrots (Psittacula krameri). Vet Microbiol. 2019;239:108428. [DOI] [PubMed] [Google Scholar]
85.Ackermann M, Koriabine M, Hartmann-Fritsch F, de Jong PJ, Lewis TD, Schetle N, et al. The genome of chelonid herpesvirus 5 harbors atypical genes. PLoS One. 2012;7(10):e46623. [DOI] [PMC free article] [PubMed] [Google Scholar]
86.Gandar F, Wilkie GS, Gatherer D, Kerr K, Marlier D, Diez M, et al. The Genome of a tortoise herpesvirus (Testudinid Herpesvirus 3) has a novel structure and contains a large region that is not required for replication in vitro or virulence in vivo. J Virol. 2015;89(22):11438–56. [DOI] [PMC free article] [PubMed] [Google Scholar]
87.Dolan A, Cunningham C, Hector RD, Hassan-Walker AF, Lee L, Addison C, et al. Genetic content of wild-type human cytomegalovirus. J Gen Virol. 2004;85(Pt 5):1301–12. [DOI] [PubMed] [Google Scholar]
88.Hansen SG, Strelow LI, Franchi DC, Anders DG, Wong SW. Complete sequence and genomic analysis of rhesus cytomegalovirus. J Virol. 2003;77(12):6620–36. [DOI] [PMC free article] [PubMed] [Google Scholar]
89.Marsh AK, Willer DO, Ambagala APN, Dzamba M, Chan JK, Pilon R, et al. Genomic sequencing and characterization of cynomolgus macaque cytomegalovirus. J Virol. 2011;85(24):12995–3009. [DOI] [PMC free article] [PubMed] [Google Scholar]
90.Blewett EL, Sherrod CJ, Texier JR, Conrad TM, Dittmer DP. Complete genome sequences of mandrillus leucophaeus and papio ursinus cytomegaloviruses. Genome Announc. 2015;3(4):e00781. 10.1128/genomea.00781-15. [DOI] [PMC free article] [PubMed] [Google Scholar]
91.Davison AJ, Dolan A, Akter P, Addison C, Dargan DJ, Alcendor DJ, et al. The human cytomegalovirus genome revisited: comparison with the chimpanzee cytomegalovirus genome. J Gen Virol. 2003;84(Pt 1):17–28. [DOI] [PubMed] [Google Scholar]
92.Cheng TP, Valentine MC, Gao J, Pingel JT, Yokoyama WM. Stability of murine cytomegalovirus genome after in vitro and in vivo passage. J Virol. 2010;84(5):2623–8. [DOI] [PMC free article] [PubMed] [Google Scholar]
93.Vink C, Beuken E, Bruggeman CA. Complete DNA sequence of the rat cytomegalovirus genome. J Virol. 2000;74(16):7656–65. [DOI] [PMC free article] [PubMed] [Google Scholar]
94.Ettinger J, Geyer H, Nitsche A, Zimmermann A, Brune W, Sandford GR, et al. Complete genome sequence of the english isolate of rat cytomegalovirus (Murid Herpesvirus 8). J Virol. 2012;86(24):13838–13838. [DOI] [PMC free article] [PubMed] [Google Scholar]
95.Yang D, Tamburro K, Dittmer D, Cui X, McVoy MA, Hernandez-Alvarado N, et al. Complete genome sequence of pathogenic guinea pig cytomegalovirus from salivary gland homogenates of infected animals. Genome Announc. 2013;1(2):e0005413. 10.1128/genomea.00054-13. [DOI] [PMC free article] [PubMed] [Google Scholar]
96.Zhang H, Todd S, Tachedjian M, Barr JA, Luo M, Yu M, et al. A novel bat herpesvirus encodes homologues of major histocompatibility complex classes I and II, C-type lectin, and a unique family of immune-related genes. J Virol. 2012;86(15):8014–30. [DOI] [PMC free article] [PubMed] [Google Scholar]
97.Bahr U, Darai G. Analysis and characterization of the complete genome of tupaia (tree shrew) herpesvirus. J Virol. 2001;75(10):4854–70. [DOI] [PMC free article] [PubMed] [Google Scholar]
98.Megaw AG, Rapaport D, Avidor B, Frenkel N, Davison AJ. The DNA sequence of the RK strain of human herpesvirus 7. Virology. 1998;244(1):119–32. [DOI] [PubMed] [Google Scholar]
99.Gompels UA, Nicholas J, Lawrence G, Jones M, Thomson BJ, Martin ME, et al. The DNA sequence of human herpesvirus-6: structure, coding content, and genome evolution. Virology. 1995;209(1):29–51. [DOI] [PubMed] [Google Scholar]
100.Dominguez G, Dambaugh TR, Stamey FR, Dewhurst S, Inoue N, Pellett PE. Human herpesvirus 6B genome sequence: coding content and comparison with human herpesvirus 6A. J Virol. 1999;73(10):8040–52. [DOI] [PMC free article] [PubMed] [Google Scholar]
101.Staheli JP, Dyen MR, Deutsch GH, Basom RS, Fitzgibbon MP, Lewis P, et al. Complete unique genome sequence, expression profile, and salivary gland tissue tropism of the herpesvirus 7 homolog in pigtailed macaques. J Virol. 2016;90(15):6657–74. [DOI] [PMC free article] [PubMed] [Google Scholar]
102.Patel SJ, Zhao G, Penna VR, Park E, Lauron EJ, Harvey IB, et al. A Murine Herpesvirus Closely Related to Ubiquitous Human Herpesviruses Causes T-Cell Depletion. J Virol. 2017;91(9):e02463-16 Sandri-Goldin RM, éditeur. [DOI] [PMC free article] [PubMed] [Google Scholar]
103.Wilkie GS, Davison AJ, Watson M, Kerr K, Sanderson S, Bouts T, et al. Complete genome sequences of elephant endotheliotropic herpesviruses 1A and 1B determined directly from fatal cases. J Virol. 2013;87(12):6700–12. [DOI] [PMC free article] [PubMed] [Google Scholar]
104.Long SY, Latimer EM, Hayward GS. Review of elephant endotheliotropic herpesviruses and acute hemorrhagic disease. ILAR J. 2016;56(3):283–96. [DOI] [PMC free article] [PubMed] [Google Scholar]
105.Ling PD, Long SY, Fuery A, Peng RS, Heaggans SY, Qin X, et al. Complete Genome Sequence of Elephant Endotheliotropic Herpesvirus 4, the First Example of a GC-Rich Branch Proboscivirus. mSphere. 2016;1(3):e00081-15. [DOI] [PMC free article] [PubMed] [Google Scholar]
106.Davison AJ, Subramaniam K, Kerr K, Jacob JM, Landrau-Giovannetti N, Walsh MT, et al. Genome Sequence of a Gammaherpesvirus from a Common Bottlenose Dolphin (Tursiops truncatus). Genome Announc. 2017;5(31):e00777-e817. [DOI] [PMC free article] [PubMed] [Google Scholar]
107.Ensser A, Pflanz R, Fleckenstein B. Primary structure of the alcelaphine herpesvirus 1 genome. J Virol. 1997;71(9):6517–25. [DOI] [PMC free article] [PubMed] [Google Scholar]
108.Taus NS, O’Toole D, Herndon DR, Cunha CW, Warg JV, Seal BS, et al. Malignant catarrhal fever in American bison (Bison bison) experimentally infected with alcelaphine herpesvirus 2. Vet Microbiol. 2014;172(1):318–22. [DOI] [PubMed] [Google Scholar]
109.Jia J, Delhon G, Tulman ER, Diel DG, Osorio FA, Wen X, et al. Novel gammaherpesvirus functions encoded by bovine herpesvirus 6 (bovine lymphotropic virus). J Gen Virol. 2014;95(8):1790–8. [DOI] [PubMed] [Google Scholar]
110.Chmielewicz B, Goltz M, Ehlers B. Detection and multigenic characterization of a novel gammaherpesvirus in goats. Virus Res. 2001;75(1):87–94. [DOI] [PubMed] [Google Scholar]
111.Li H, Gailbreath K, Flach EJ, Taus NS, Cooley J, Keller J, et al. A novel subgroup of rhadinoviruses in ruminants. J Gen Virol. 2005;86(Pt 11):3021–6. [DOI] [PubMed] [Google Scholar]
112.Hart J, Ackermann M, Jayawardane G, Russell G, Haig DM, Reid H, et al. Complete sequence and analysis of the ovine herpesvirus 2 genome. J Gen Virol. 2007;88(1):28–39. [DOI] [PubMed] [Google Scholar]
113.Ehlers B, Ulrich S, Goltz M. Detection of two novel porcine herpesviruses with high similarity to gammaherpesviruses. J Gen Virol. 1999;80(4):971–8. [DOI] [PubMed] [Google Scholar]
114.Chmielewicz B, Goltz M, Franz T, Bauer C, Brema S, Ellerbrok H, et al. A novel porcine gammaherpesvirus. Virology. 2003;308(2):317–29. [DOI] [PubMed] [Google Scholar]
115.Telford EAR, Watson MS, Aird HC, Perry J, Davison AJ. The DNA sequence of equine herpesvirus 2. J Mol Biol. 1995;249(3):520–8. [DOI] [PubMed] [Google Scholar]
116.Wilkie GS, Kerr K, Stewart JP, Studdert MJ, Davison AJ. Genome Sequences of equid herpesviruses 2 and 5. Genome Announce. 2015;3(2):e00119. 10.1128/genomea.00119-15. [DOI] [PMC free article] [PubMed] [Google Scholar]
117.Troyer RM, Lee JS, Vuyisich M, Chain P, Lo CC, Kronmiller B, et al. First complete genome sequence of felis catus gammaherpesvirus 1. Genome Announc. 2015;3(6):e01192. 10.1128/genomea.01192-15. [DOI] [PMC free article] [PubMed] [Google Scholar]
118.Banks M, King DP, Daniells C, Stagg DA, Gavier-Widen D. Partial characterization of a novel gammaherpesvirus isolated from a European badger (Meles meles). J Gen Virol. 2002;83(Pt 6):1325–30. [DOI] [PubMed] [Google Scholar]
119.Bellehumeur C, Nielsen O, Measures L, Harwood L, Goldstein T, Boyle B, et al. Herpesviruses including novel gammaherpesviruses are widespread among phocid seal species in Canada. J Wildl Dis. 2016;52(1):70–81. [DOI] [PubMed] [Google Scholar]
120.Noguchi K, Kuwata R, Shimoda H, Mizutani T, Hondo E, Maeda K. The complete genomic sequence of Rhinolophus gammaherpesvirus 1 isolated from a greater horseshoe bat. Arch Virol. 2019;164(1):317–9. [DOI] [PubMed] [Google Scholar]
121.Shabman RS, Shrivastava S, Tsibane T, Attie O, Jayaprakash A, Mire CE, et al. Isolation and characterization of a novel gammaherpesvirus from a microbat cell line. mSphere. 2016;1(1):e00070-15. [DOI] [PMC free article] [PubMed] [Google Scholar]
122.Richter J, Puchtler I, Fleckenstein B. Thymidylate synthase gene of herpesvirus ateles. J Virol. 1988;62(9):3530–5. [DOI] [PMC free article] [PubMed] [Google Scholar]
123.Albrecht JC. Primary structure of the Herpesvirus ateles genome. J Virol. 2000;74(2):1033–7. [DOI] [PMC free article] [PubMed] [Google Scholar]
124.Zimmermann W, Broll H, Ehlers B, Buhk HJ, Rosenthal A, Goltz M. Genome sequence of bovine herpesvirus 4, a bovine Rhadinovirus, and identification of an origin of DNA replication. J Virol. 2001;75(3):1186–94. [DOI] [PMC free article] [PubMed] [Google Scholar]
125.Dhingra A, Ganzenmueller T, Hage E, Suárez NM, Mätz-Rensing K, Widmer D, et al. Novel Virus Related to Kaposi’s Sarcoma–Associated Herpesvirus from Colobus Monkey - Volume 25, Number 8—August 2019 - Emerging Infectious Diseases journal - CDC. 2019; Disponible sur: https://wwwnc.cdc.gov/eid/article/25/8/18-1802_article. Cité 16 Juill 2024. [DOI] [PMC free article] [PubMed]
126.Loh J, Zhao G, Nelson CA, Coder P, Droit L, Handley SA, et al. Identification and sequencing of a novel rodent gammaherpesvirus that establishes acute and latent infection in laboratory mice. J Virol. 2011;85(6):2642–56. [DOI] [PMC free article] [PubMed] [Google Scholar]
127.Glenn M, Rainbow L, Auradé F, Davison A, Schulz TF. Identification of a spliced gene from Kaposi’s sarcoma-associated herpesvirus encoding a protein with similarities to latent membrane proteins 1 and 2A of Epstein-Barr virus. J Virol. 1999;73(8):6953–63. [DOI] [PMC free article] [PubMed] [Google Scholar]
128.Axthelm MK, Bourdette DN, Marracci GH, Su W, Mullaney ET, Manoharan M, et al. Japanese macaque encephalomyelitis: a spontaneous multiple sclerosis–like disease in a nonhuman primate. Ann Neurol. 2011;70(3):362–73. [DOI] [PMC free article] [PubMed] [Google Scholar]
129.Bruce AG, Thouless ME, Haines AS, Pallen MJ, Grundhoff A, Rose TM. Complete genome sequence of Pig-tailed macaque rhadinovirus 2 and its evolutionary relationship with rhesus macaque rhadinovirus and human herpesvirus 8/Kaposi’s sarcoma-associated herpesvirus. J Virol. 2015;89(7):3888–909. [DOI] [PMC free article] [PubMed] [Google Scholar]
130.Searles RP, Bergquam EP, Axthelm MK, Wong SW. Sequence and Genomic Analysis of a Rhesus Macaque Rhadinovirus with Similarity to Kaposi’s Sarcoma-Associated Herpesvirus/Human Herpesvirus 8. J Virol. 1999;73(4):3040–53. [DOI] [PMC free article] [PubMed] [Google Scholar]
131.Bruce AG, Ryan JT, Thomas MJ, Peng X, Grundhoff A, Tsai CC, et al. Next-generation sequence analysis of the genome of RFHVMn, the macaque homolog of Kaposi’s sarcoma (KS)-associated herpesvirus, from a KS-like tumor of a pig-tailed macaque. J Virol. 2013;87(24):13676–93. [DOI] [PMC free article] [PubMed] [Google Scholar]
132.Virgin HW, Latreille P, Wamsley P, Hallsworth K, Weck KE, Dal Canto AJ, et al. Complete sequence and genomic analysis of murine gammaherpesvirus 68. J Virol. 1997;71(8):5894–904. [DOI] [PMC free article] [PubMed] [Google Scholar]
133.Hughes DJ, Kipar A, Milligan SG, Cunningham C, Sanders M, Quail MA, et al. Characterization of a novel wood mouse virus related to murid herpesvirus 4. J Gen Virol. 2010;91(Pt 4):867–79. [DOI] [PMC free article] [PubMed] [Google Scholar]
134.Albrecht JC, Nicholas J, Biller D, Cameron KR, Biesinger B, Newman C, et al. Primary structure of the herpesvirus saimiri genome. J Virol. 1992;66(8):5047–58. [DOI] [PMC free article] [PubMed] [Google Scholar]
135.Subudhi S, Rapin N, Dorville N, Hill JE, Town J, Willis CKR, et al. Isolation, characterization and prevalence of a novel Gammaherpesvirus in Eptesicus fuscus, the North American big brown bat. Virology. 2018;516:227–38. [DOI] [PubMed] [Google Scholar]
136.Vaz PK, Hartley CA, Lee SY, Sansom FM, Adams TE, Stalder K, et al. Koala and Wombat Gammaherpesviruses Encode the First Known Viral NTPDase Homologs and Are Phylogenetically Divergent from All Known Gammaherpesviruses. J Virol. 2019;93(6):e01404-e1418. [DOI] [PMC free article] [PubMed] [Google Scholar]
137.Cho Y, Ramer J, Rivailler P, Quink C, Garber RL, Beier DR, et al. An Epstein-Barr-related herpesvirus from marmoset lymphomas. Proc Natl Acad Sci U S A. 2001;98(3):1224–9. [DOI] [PMC free article] [PubMed] [Google Scholar]
138.Gerner CS, Dolan A, McGeoch DJ. Phylogenetic relationships in the Lymphocryptovirus genus of the Gammaherpesvirinae. Virus Res. 2004;99(2):187–92. [DOI] [PubMed] [Google Scholar]
139.Baer R, Bankier AT, Biggin MD, Deininger PL, Farrell PJ, Gibson TJ, et al. DNA sequence and expression of the B95–8 Epstein—Barr virus genome. Nature. 1984;310(5974):207–11. [DOI] [PubMed] [Google Scholar]
140.Kamperschroer C, Gosink MM, Kumpf SW, O’Donnell LM, Tartaro KR. The genomic sequence of lymphocryptovirus from cynomolgus macaque. Virology. 2016;488:28–36. [DOI] [PubMed] [Google Scholar]
141.Krumbholz A, Roempke J, Liehr T, Groth M, Meerbach A, Schacke M, et al. Macaca arctoides gammaherpesvirus 1 (strain herpesvirus Macaca arctoides): virus sequence, phylogeny and characterisation of virus-transformed macaque and rabbit cell lines. Med Microbiol Immunol. 2019;208(1):109–29. [DOI] [PubMed] [Google Scholar]
142.Rivailler P, Jiang H, Cho YG, Quink C, Wang F. Complete Nucleotide Sequence of the Rhesus Lymphocryptovirus: Genetic Validation for an Epstein-Barr Virus Animal Model. J Virol. 2002;76(1):421–6. [DOI] [PMC free article] [PubMed] [Google Scholar]
143.Savin KW, Cocks BG, Wong F, Sawbridge T, Cogan N, Savage D, et al. A neurotropic herpesvirus infecting the gastropod, abalone, shares ancestry with oyster herpesvirus and a herpesvirus associated with the amphioxus genome. Virol J. 2010;7(1):308. [DOI] [PMC free article] [PubMed] [Google Scholar]
144.Davison AJ, Cunningham C, Sauerbier W, McKinnell RG. Genome sequences of two frog herpesviruses. J Gen Virol. 2006;87(12):3509–14. [DOI] [PubMed] [Google Scholar]
145.Origgi FC, Schmidt BR, Lohmann P, Otten P, Akdesir E, Gaschen V, et al. Ranid herpesvirus 3 and proliferative dermatitis in free-ranging wild common frogs (Rana Temporaria). Vet Pathol. 2017;54(4):686–94. [DOI] [PubMed] [Google Scholar]
146.Doszpoly A, Kovács ER, Bovo G, LaPatra SE, Harrach B, Benko M. Molecular confirmation of a new herpesvirus from catfish (Ameiurus melas) by testing the performance of a novel PCR method, designed to target the DNA polymerase gene of alloherpesviruses. Arch Virol. 2008;153(11):2123–7. [DOI] [PubMed] [Google Scholar]
147.Davison AJ. Channel catfish virus: a new type of herpesvirus. Virology. 1992;186(1):9–14. [DOI] [PubMed] [Google Scholar]
148.Borzák R, Haluk T, Bartha D, Doszpoly A. Complete genome sequence and analysis of ictalurid herpesvirus 2. Arch Virol. 2018;163(4):1083–5. [DOI] [PubMed] [Google Scholar]
149.van Beurden SJ, Bossers A, Voorbergen-Laarman MHA, Haenen OLM, Peters S, Abma-Henkens MHC, et al. Complete genome sequence and taxonomic position of anguillid herpesvirus 1. J Gen Virol. 2010;91(4):880–7. [DOI] [PubMed] [Google Scholar]
150.Davison AJ, Kurobe T, Gatherer D, Cunningham C, Korf I, Fukuda H, et al. Comparative genomics of carp herpesviruses. J Virol. 2013;87(5):2908–22. [DOI] [PMC free article] [PubMed] [Google Scholar]
151.Aoki T, Hirono I, Kurokawa K, Fukuda H, Nahary R, Eldar A, et al. Genome sequences of three koi herpesvirus isolates representing the expanding distribution of an emerging disease threatening koi and common carp worldwide. J Virol. 2007;81(10):5058–65. [DOI] [PMC free article] [PubMed] [Google Scholar]
152.Davison AJ. The genome of salmonid herpesvirus 1. J Virol. 1998;72(3):1974–82. [DOI] [PMC free article] [PubMed] [Google Scholar]
153.Waltzek T, Kelley G, Alfaro M, Kurobe T, Davison A, Hedrick R. Phylogenetic relationships in the family Alloherpesviridae. Dis Aquat Org. 2009;84:179–94. [DOI] [PubMed] [Google Scholar]
154.Boutier M, Morvan L, Delrez N, Origgi F, Doszpoly A, Vanderplasschen A. Fish and Amphibian Alloherpesviruses (Herpesviridae). In: Encyclopedia of Virology. Elsevier; 2021. p. 306‑15. Disponible sur: https://linkinghub.elsevier.com/retrieve/pii/B978012809633820931X. Cité 8 août 2023.
155.Rosani U, Gaia M, Delmont TO, Krupovic M. Tracing the invertebrate herpesviruses in the global sequence datasets. Front Mar Sci. 2023;10:1159754. [Google Scholar]
156.Ferreira RC, Alves GV, Ramon M, Antoneli F, Briones MRS. Reconstructing prehistoric viral genomes from neanderthal sequencing data. Viruses. 2024;16(6):856. [DOI] [PMC free article] [PubMed] [Google Scholar]
157.Alfano N, Dayaram A, Axtner J, Tsangaras K, Kampmann ML, Mohamed A, et al. Non-invasive surveys of mammalian viruses using environmental DNA. Methods Ecol Evol. 2021;12(10):1941–52. [Google Scholar]
158.Bass D, Christison KW, Stentiford GD, Cook LSJ, Hartikainen H. Environmental DNA/RNA for pathogen and parasite detection, surveillance, and ecology. Trends Parasitol. 2023;39(4):285–304. [DOI] [PubMed] [Google Scholar]
159.Bavarsad M, Abed-Elmdoost A, Tabandeh MR, Alishahi M, Mirvaghefi A, Farahmand H. Rapid cyprinid herpesvirus 3 detection using loop-mediated isothermal amplification (LAMP) combined with gold nanoprobes and SYBR safe. Aquaculture. 2025;594:741374. [Google Scholar]
160.Dotto-Maurel A, Pelletier C, Morga B, Jacquot M, Faury N, Dégremont L, et al. Evaluation of tangential flow filtration coupled to long-read sequencing for ostreid herpesvirus type 1 genome assembly. Microbial Genomics. 2022;8(11). Disponible sur: https://www.microbiologyresearch.org/content/journal/mgen/. 10.1099/mgen.0.000895. Cité 14 Nov 2022. [DOI] [PMC free article] [PubMed]
161.Hanson L, Dishon A, Kotler M. Herpesviruses that infect fish. Viruses. 2011;3(11):2160–91. [DOI] [PMC free article] [PubMed] [Google Scholar]
162.Khanthaphixay B, Wu L, Yoon JY. Microparticle-based detection of viruses. Biosensors. 2023;13(8):820. [DOI] [PMC free article] [PubMed] [Google Scholar]
163.Volpe E, Errani F, Mandrioli L, Ciulli S. Advances in viral aquatic animal disease knowledge: the molecular methods’ contribution. Biology mars. 2023;12(3):466. [DOI] [PMC free article] [PubMed] [Google Scholar]
164.Gu W, Zeng N, Zhou L, Ge X, Guo X, Yang H. Genomic organization and molecular characterization of porcine cytomegalovirus. Virology. 2014;460–461:165–72. [DOI] [PubMed] [Google Scholar]
165.Chang W, Jiao X, Sui H, Goswami S, Sherman BT, Fromont C, et al. Complete genome sequence of herpes simplex virus 2 strain G. Viruses. 2022;14(3):536. [DOI] [PMC free article] [PubMed] [Google Scholar]
166.Delius H, Clements JB. A partial denaturation map of herpes simplex virus Type 1 DNA: evidence for inversions of the unique DNA Regions. J Gen Virol. 1976;33(1):125–33. [DOI] [PubMed] [Google Scholar]
167.Desrosiers RC, Falk LA. Herpesvirus tamarinus and its relation to herpes simplex virus. J Gen Virol. 1981;56(1):119–30. [DOI] [PubMed] [Google Scholar]
168.Henry BE, Robinson RA, Dauenhauer SA, Atherton SS, Hayward GS, O’Callaghan DJ. Structure of the genome of equine herpesvirus type 1. Virology. 1981;115(1):97–114. [DOI] [PubMed] [Google Scholar]
169.Jenkins FJ, Roizman B. Herpes simplex virus 1 recombinants with noninverting genomes frozen in different isomeric arrangements are capable of independent replication. J Virol. 1986;59(2):494–9. [DOI] [PMC free article] [PubMed] [Google Scholar]
170.Leib DA, Bradbury JM, Hart CA, McCarthy K. Genome isomerism in two alphaherpesviruses: Herpesvirus saimiri-1(Herpesvirus tamarinus) and avian infectious laryngotracheitis virus. Adv Virol. 1987;93(3):287–94. [DOI] [PubMed] [Google Scholar]
171.Lomniczi B, Blankenship ML, Ben-Porat T. Deletions in the genomes of pseudorabies virus vaccine strains and existence of four isomers of the genomes. J Virol. 1984;49(3):970–9. [DOI] [PMC free article] [PubMed] [Google Scholar]
172.Sauer A, Wang JB, Hahn G, McVoy MA. A human cytomegalovirus deleted of internal repeats replicates with near wild type efficiency but fails to undergo genome isomerization. Virology. 2010;401(1):90–5. [DOI] [PMC free article] [PubMed] [Google Scholar]
173.Schynts F, McVoy MA, Meurens F, Detry B, Epstein AL, Thiry E. The structures of bovine herpesvirus 1 virion and concatemeric DNA: implications for cleavage and packaging of herpesvirus genomes. Virology. 2003;314(1):326–35. [DOI] [PubMed] [Google Scholar]
174.Wood ML, Neumann R, Roy P, Nair V, Royle NJ. Characterization of integrated Marek’s disease virus genomes supports a model of integration by homology-directed recombination and telomere-loop-driven excision. J Virol. 2023;97(10):e00716-e723. [DOI] [PMC free article] [PubMed] [Google Scholar]
175.Mahiet C, Ergani A, Huot N, Alende N, Azough A, Salvaire F, et al. Structural variability of the herpes simplex virus 1 genome in vitro and in vivo. J Virol. 2012;86(16):8592–601. [DOI] [PMC free article] [PubMed] [Google Scholar]
176.Kinchington PR, Reinhold WC, Casey TA, Straus SE, Hay J, Ruyechan WT. Inversion and circularization of the varicella-zoster virus genome. J Virol. 1985;56(1):194–200. [DOI] [PMC free article] [PubMed] [Google Scholar]
177.Nicolson L. The Characterisation of Specific EHV-4 Genes [PhD]. [Ann Arbor :]: ProQuest Dissertations & Theses; 1991. Disponible sur: https://theses.gla.ac.uk/77027/. Cité 11 Sept 2023.
178.Weber PC, Challberg MD, Nelson NJ, Levine M, Glorioso JC. Inversion events in the HSV-1 genome are directly mediated by the viral DNA replication machinery and lack sequence specificity. Cell. 1988;54(3):369–81. [DOI] [PubMed] [Google Scholar]
179.Jacob RJ, Morse LS, Roizman B. Anatomy of herpes simplex virus DNA XII. Accumulation of head-to-tail concatemers in nuclei of infected cells and their role in the generation of the four isomeric arrangements of viral DNA. J Virol. 1979;29(2):448–57. [DOI] [PMC free article] [PubMed] [Google Scholar]
180.Roizman B. The structure and isomerization of herpes simplex virus genomes. Cell. 1979;16(3):481–94. [DOI] [PubMed] [Google Scholar]
181.Hayward GS, Jacob RJ, Wadsworth SC, Roizman B. Anatomy of herpes simplex virus DNA: evidence for four populations of molecules that differ in the relative orientations of their long and short components. 1975. Disponible sur: https://www.pnas.org/doi/10.1073/pnas.72.11.4243. Cité 3 Sept 2023. [DOI] [PMC free article] [PubMed]
182.McKenzie J, El-Guindy A. Epstein-Barr Virus Lytic Cycle Reactivation. In: Münz C, éditeur. Epstein Barr Virus Volume 2: One Herpes Virus: Many Diseases. Cham: Springer International Publishing; 2015. p. 237‑61. (Current Topics in Microbiology and Immunology). Disponible sur: 10.1007/978-3-319-22834-1_8. Cité 17 juill 2023. [DOI] [PubMed]
183.Boldogh I, Albrecht T, Porter DD. Persistent Viral Infections. In: Baron S, éditeur. Medical Microbiology. 4th éd. Galveston (TX): University of Texas Medical Branch at Galveston; 1996. Disponible sur: http://www.ncbi.nlm.nih.gov/books/NBK8538/. Cité 27 Juill 2023. [PubMed]
184.Brunner S, Herndler-Brandstetter D, Weinberger B, Grubeck-Loebenstein B. Persistent viral infections and immune aging. Ageing Res Rev. 2011;10(3):362–9. [DOI] [PubMed] [Google Scholar]
185.Frebel H, Richter K, Oxenius A. How chronic viral infections impact on antigen-specific T-cell responses. Eur J Immunol. 2010;40(3):654–63. [DOI] [PubMed] [Google Scholar]
186.Gerna G, Kabanova A, Lilleri D. Human cytomegalovirus cell tropism and host cell receptors. Vaccines (Basel). 2019;7(3):70. [DOI] [PMC free article] [PubMed] [Google Scholar]
187.Wofford AS, McCusker I, Green JC, Vensko TA, Pellett PE. Chapter Ten - Betaherpesvirus assembly and egress: Recent advances illuminate the path. In: Kielian M, Mettenleiter TC, Roossinck MJ, éditeurs. Advances in Virus Research. Academic Press; 2020. p. 337‑92. (Virus Assembly and Exit Pathways; vol. 108). Disponible sur: https://www.sciencedirect.com/science/article/pii/S0065352720300439. Cité 25 Nov 2024. [DOI] [PubMed]
188.Liashkovich I, Hafezi W, Kühn JM, Oberleithner H, Shahin V. Nuclear delivery mechanism of herpes simplex virus type 1 genome. J Mol Recognit. 2011;24(3):414–21. [DOI] [PubMed] [Google Scholar]
189.Kukhanova MK, Korovina AN, Kochetkov SN. Human herpes simplex virus: Life cycle and development of inhibitors. Biochemistry Moscow. 2014;79(13):1635–52. [DOI] [PubMed] [Google Scholar]
190.Castillo JP, Kowalik TF. HCMV infection: modulating the cell cycle and cell death. Int Rev Immunol. 2004;23(1–2):113–39. [DOI] [PubMed] [Google Scholar]
191.Damania B, Kenney SC, Raab-Traub N. Epstein-Barr virus: biology and clinical disease. Cell. 2022;185(20):3652–70. [DOI] [PMC free article] [PubMed] [Google Scholar]
192.Boehmer P, Nimonkar A. Herpes virus replication. IUBMB Life. 2003;55(1):13–22. [DOI] [PubMed] [Google Scholar]
193.Zhu S, Viejo-Borbolla A. Pathogenesis and virulence of herpes simplex virus. Virulence. 2021;12(1):2670–702. [DOI] [PMC free article] [PubMed] [Google Scholar]
194.DeLuca NA, Schaffer PA. Activation of immediate-early, early, and late promoters by temperature-sensitive and wild-type forms of herpes simplex virus type 1 protein ICP4. Mol Cell Biol. 1985;5(8):1997–2008. [DOI] [PMC free article] [PubMed] [Google Scholar]
195.Hardwicke MA, Sandri-Goldin RM. The herpes simplex virus regulatory protein ICP27 contributes to the decrease in cellular mRNA levels during infection. J Virol. 1994;68(8):4797–810. [DOI] [PMC free article] [PubMed] [Google Scholar]
196.Spencer CA, Dahmus ME, Rice SA. Repression of host RNA polymerase II transcription by herpes simplex virus type 1. J Virol. 1997;71(3):2031–40. [DOI] [PMC free article] [PubMed] [Google Scholar]
197.Homa FL, Brown JC. Capsid assembly and DNA packaging in herpes simplex virus. Rev Med Virol. 1997;7(2):107–22. [DOI] [PubMed] [Google Scholar]
198.Close WL, Anderson AN, Pellett PE. Betaherpesvirus Virion Assembly and Egress. In: Kawaguchi Y, Mori Y, Kimura H, éditeurs. Human Herpesviruses. Singapore: Springer; 2018. p. 167‑207. Disponible sur: 10.1007/978-981-10-7230-7_9. Cité 17 Nov 2024.
199.Crump C. Virus assembly and egress of HSV. Adv Exp Med Biol. 2018;1045:23–44. [DOI] [PubMed] [Google Scholar]
200.Hellberg T, Paßvogel L, Schulz KS, Klupp BG, Mettenleiter TC. Nuclear egress of herpesviruses: the prototypic vesicular nucleocytoplasmic transport. Adv Virus Res. 2016;94:81–140. [DOI] [PubMed] [Google Scholar]
201.Roller RJ, Baines JD. Herpesvirus nuclear egress. Adv Anat Embryol Cell Biol. 2017;223:143–69. [DOI] [PubMed] [Google Scholar]
202.Dodding MP, Way M. Coupling viruses to dynein and kinesin‐1. The EMBO Journal. 2011; Disponible sur: https://www.embopress.org/doi/10.1038/emboj.2011.283. Cité 25 2024. [DOI] [PMC free article] [PubMed]
203.Lyman MG, Enquist LW. Herpesvirus interactions with the host cytoskeleton. J Virol. 2009;83(5):2058–66. [DOI] [PMC free article] [PubMed] [Google Scholar]
204.Naghavi MH, Walsh D. Microtubule regulation and function during virus infection. J Virol. 2017;91(16):e00538-e617. [DOI] [PMC free article] [PubMed] [Google Scholar]
205.Abdelgawad A, Damiani A, Ho SYW, Strauss G, Szentiks CA, East ML, et al. Zebra Alphaherpesviruses (EHV-1 and EHV-9): genetic diversity, latency and co-infections. Viruses. 2016;8(9):262. [DOI] [PMC free article] [PubMed] [Google Scholar]
206.Aghazadeh M, Shi M, Pesavento PA, Durham AC, Polley T, Donahoe SL, et al. Transcriptome analysis and in situ hybridization for FcaGHV1 in feline lymphoma. Viruses. 2018;10(9):464. [DOI] [PMC free article] [PubMed] [Google Scholar]
207.Aligo J, Walker M, Bugelski P, Weinstock D. Is murine gammaherpesvirus-68 (MHV-68) a suitable immunotoxicological model for examining immunomodulatory drug-associated viral recrudescence? J Immunotoxicol. 2015;12(1):1–15. [DOI] [PubMed] [Google Scholar]
208.Bielefeldt-Ohmann H, Bruce AG, Howard K, Ikoma M, Thouless ME, Rose TM. Macaque homologs of Kaposi’s sarcoma-associated herpesvirus (KSHV) infect germinal center lymphoid cells, epithelial cells in skin and gastrointestinal tract and gonadal germ cells in naturally infected macaques. Virology. 2018;519:106–20. [DOI] [PMC free article] [PubMed] [Google Scholar]
209.Broussard G, Damania B. Regulation of KSHV latency and lytic reactivation. Viruses. 2020;12(9):1034. [DOI] [PMC free article] [PubMed] [Google Scholar]
210.Camero M, Lanave G, Lucente MS, Losurdo M, Di Paola G, Lorusso E, et al. Bubaline alphaherpesvirus 1 induces a latent/reactivable infection in goats. Comp Immunol Microbiol Infect Dis. 2019;62:54–7. [DOI] [PubMed] [Google Scholar]
211.Campos FS, Franco AC, Oliveira MT, Firpo R, Strelczuk G, Fontoura FE, et al. Detection of bovine herpesvirus 2 and bovine herpesvirus 4 DNA in trigeminal ganglia of naturally infected cattle by polymerase chain reaction. Vet Microbiol. 2014;171(1):182–8. [DOI] [PubMed] [Google Scholar]
212.Collins CM, Medveczky MM, Lund T, Medveczky PG. The terminal repeats and latency-associated nuclear antigen of herpesvirus saimiri are essential for episomal persistence of the viral genome. J Gen Virol. 2002;83(9):2269–78. [DOI] [PubMed] [Google Scholar]
213.Darai G, Koch HG. Tree Shrew Herpesvirus: Pathogenicity and Latency. In: Wittmann G, Gaskell RM, Rziha HJ, éditeurs. Latent Herpes Virus Infections in Veterinary Medicine: A Seminar in the CEC Programme of Coordination of Research on Animal Pathology, held at Tübingen, Federal Republic of Germany, September 21–24, 1982. Dordrecht: Springer Netherlands; 1984. p. 91‑102. Disponible sur: 10.1007/978-94-009-5662-9_8. Cité 19 févr 2025.
214.das Neves CG, Mørk T, Thiry J, Godfroid J, Rimstad E, Thiry E, et al. Cervid herpesvirus 2 experimentally reactivated in reindeer can produce generalized viremia and abortion. Virus Res. 2009;145(2):321–8. [DOI] [PubMed] [Google Scholar]
215.Denesvre C, You Y, Rémy S, Vychodil T, Courvoisier K, Penzes Z, et al. Impact of viral telomeric repeat sequences on herpesvirus vector vaccine integration and persistence. PLOS Pathogens. 2024;20(5):e1012261. [DOI] [PMC free article] [PubMed] [Google Scholar]
216.Dewals BG, Vanderplasschen A. Malignant catarrhal fever induced by Alcelaphine herpesvirus 1 is characterized by an expansion of activated CD3+CD8+CD4- T cells expressing a cytotoxic phenotype in both lymphoid and non-lymphoid tissues. Vet Res. 2011;42(1):95. [DOI] [PMC free article] [PubMed] [Google Scholar]
217.Divilov K, Wang X, Swisher AE, Yeoman PC, Rintoul M, Fleener GB, et al. Ostreid herpesvirus 1 latent infection and reactivation in adult Pacific oysters, Crassostrea gigas. Virus Res. 2024;339:199245. [DOI] [PMC free article] [PubMed] [Google Scholar]
218.Eberle R, Maxwell LK, Nicholson S, Black D, Jones-Engel L. Genome sequence variation among isolates of monkey B virus (Macacine alphaherpesvirus 1) from captive macaques. Virology. 2017;508:26–35. [DOI] [PMC free article] [PubMed] [Google Scholar]
219.Freifeld AG, Hilliard J, Southers J, Murray M, Savarese B, Schmitt JM, et al. A controlled seroprevalence survey of primate handlers for evidence of asymptomatic herpes B virus infection. J Infect Dis. 1995;171(4):1031–4. [DOI] [PubMed] [Google Scholar]
220.Gaskell R, Dawson S, Radford A, Thiry E. Feline herpesvirus. Vet Res. 2007;38(2):337–54. [DOI] [PubMed] [Google Scholar]
221.Gowthaman V, Kumar S, Koul M, Dave U, Murthy TRGK, Munuswamy P, et al. Infectious laryngotracheitis: Etiology, epidemiology, pathobiology, and advances in diagnosis and control – a comprehensive review. Vet Q. 2020;40(1):140–61. [DOI] [PMC free article] [PubMed] [Google Scholar]
222.Holland MS, Mackenzie CD, Bull RW, Silva RF. Latent Turkey herpesvirus infection in lymphoid, nervous, and feather tissues of chickens. Avian Dis. 1998;42(2):292–9. [PubMed] [Google Scholar]
223.Mahalingam R, Clarke P, Wellish M, Dueland AN, Soike KF, Gilden DH, et al. Prevalence and distribution of latent simian varicella virus DNA in monkey ganglia. Virology. 1992;188(1):193–7. [DOI] [PubMed] [Google Scholar]
224.Mekuria ZH, El-Hage C, Ficorilli NP, Washington EA, Gilkerson JR, Hartley CA. Mapping B lymphocytes as major reservoirs of naturally occurring latent equine herpesvirus 5 infection. J Gen Virol. 2017;98(3):461–70. [DOI] [PubMed] [Google Scholar]
225.Mueller NJ, Denner J. Porcine cytomegalovirus/porcine roseolovirus (PCMV/PRV): a threat for xenotransplantation? Xenotransplantation. 2022;29(5):e12775. [DOI] [PubMed] [Google Scholar]
226.Pan L, Li M, Zhang X, Xia Y, Mian AM, Wu H, et al. Establishment of an in vitro model of pseudorabies virus latency and reactivation and identification of key viral latency-associated genes. Viruses. 2023;15(3):808. [DOI] [PMC free article] [PubMed] [Google Scholar]
227.Quijano Cardé EM, Soto E. A review of latency in the Alloherpesviridae family. J Fish Dis. 2024;47(12):e14016. [DOI] [PMC free article] [PubMed] [Google Scholar]
228.Riaz A, Dry I, Levy CS, Hopkins J, Grey F, Shaw DJ, et al. Ovine herpesvirus-2-encoded microRNAs target virus genes involved in virus latency. J Gen Virol. 2014;95(2):472–80. [DOI] [PMC free article] [PubMed] [Google Scholar]
229.Samoilowa S, Giessler KS, Torres CEM, Hussey GS, Allum A, Fux R, et al. Equid herpesvirus-1 distribution in equine lymphoid and neural tissues 70 days post infection. Pathogens. 2021;10(6):707. [DOI] [PMC free article] [PubMed] [Google Scholar]
230.Shawky S, Schat KA. Latency sites and reactivation of duck enteritis virus. Avian Dis. 2002;46(2):308–13. [DOI] [PubMed] [Google Scholar]
231.Silvestro C, Bratanich A. The latency related gene of bovine herpesvirus types 1 and 5 and its modulation of cellular processes. Arch Virol. 2016;161(12):3299–308. [DOI] [PubMed] [Google Scholar]
232.Thorsteinsdóttir L, Guðmundsson GÖ, Jensson H, Torsteinsdóttir S, Svansson V. Isolation of equid alphaherpesvirus 3 from a horse in Iceland with equine coital exanthema. Acta Vet Scand. 2021;63(1):6. [DOI] [PMC free article] [PubMed] [Google Scholar]
233.van Maanen C. Equine herpesvirus 1 and 4 infections: an update. Veterinary Quarterly. 2002;24(2):57–78. [PubMed] [Google Scholar]
234.Zamana RR, Gattamorta MA, Cruz Ochoa PF, Navas-Suárez PE, Sacristán C, Rossi S, et al. High Occurrence of Chelonid Alphaherpesvirus 5 (ChHV5) in Green Sea Turtles Chelonia mydas with and without Fibropapillomatosis in Feeding Areas of the São Paulo Coast, Brazil. J Aquat Anim Health. 2021;33(4):252–63. [DOI] [PubMed] [Google Scholar]
235.Jin L, Löhr CV, Vanarsdall AL, Baker RJ, Moerdyk-Schauwecker M, Levine C, et al. Characterization of a novel alphaherpesvirus associated with fatal infections of domestic rabbits. Virology. 2008;378(1):13–20. [DOI] [PubMed] [Google Scholar]
236.Slater JD, Borchers K, Thackray AM, Field HJ. The trigeminal ganglion is a location for equine herpesvirus 1 latency and reactivation in the horse. J Gen Virol. 1994;75(8):2007–16. [DOI] [PubMed] [Google Scholar]
237.Stiles J. Feline herpesvirus. Clin Tech Small Anim Pract. 2003;18(3):178–85. [DOI] [PubMed] [Google Scholar]
238.Maidana SS, Delgado F, Vagnoni L, Mauroy A, Thiry E, Romera S. Cattle are a potential reservoir of bubaline herpesvirus 1 (BuHV1). Veterinary Record Open. 2016;3(1):e000162. [DOI] [PMC free article] [PubMed] [Google Scholar]
239.Meyer G, Lemaire M, Ros C, Belak K, Gabriel A, Cassart D, et al. Comparative pathogenesis of acute and latent infections of calves with bovine herpesvirus types 1 and 5. Arch Virol. 2001;146(4):633–52. [DOI] [PubMed] [Google Scholar]
240.Ostler JB, Jones C. The bovine herpesvirus 1 latency-reactivation cycle, a chronic problem in the cattle industry. Viruses. 2023;15(2):552. [DOI] [PMC free article] [PubMed] [Google Scholar]
241.Mahalingam R, Wellish M, Wolf W, Dueland AN, Cohrs R, Vafai A, et al. Latent varicella-zoster viral DNA in human trigeminal and thoracic ganglia. N Engl J Med. 1990;323(10):627–31. [DOI] [PubMed] [Google Scholar]
242.Thilakarathne DS, Hartley CA, Diaz-Méndez A, Quinteros JA, Fakhri O, Coppo MJC, et al. Latency characteristics in specific pathogen-free chickens 21 and 35 days after intra-tracheal inoculation with vaccine or field strains of infectious laryngotracheitis virus. Avian Pathol. 2020;49(4):369–79. [DOI] [PubMed] [Google Scholar]
243.Borchers K, Wolfinger U, Lawrenz B, Schellenbach A, Ludwig H. Equine herpesvirus 4 DNA in trigeminal ganglia of naturally infected horses detected by direct in situ PCR. J Gen Virol. 1997;78(5):1109–14. [DOI] [PubMed] [Google Scholar]
244.Six A, Banks M, Engels M, Bascuñana CR, Ackermann M. Latency and reactivation of bovine herpesvirus 1 (BHV-1) in goats and of caprine herpesvirus 1 (CapHV-1) in calves. Arch Virol. 2001;146(7):1325–35. [DOI] [PubMed] [Google Scholar]
245.Borchers K, Lieckfeldt D, Ludwig A, Fukushi H, Allen G, Fyumagwa R, et al. Detection of equid herpesvirus 9 DNA in the trigeminal ganglia of a burchell’s zebra from the serengeti ecosystem. J Vet Med Sci. 2008;70(12):1377–81. [DOI] [PubMed] [Google Scholar]
246.Pantry SN, Medveczky PG. Latency, integration, and reactivation of human herpesvirus-6. Viruses. 2017;9(7):194. [DOI] [PMC free article] [PubMed] [Google Scholar]
247.Leskowitz RM, Zhou XY, Villinger F, Fogg MH, Kaur A, Lieberman PM, et al. CD4+ and CD8+ T-cell responses to latent antigen EBNA-1 and lytic antigen BZLF-1 during persistent lymphocryptovirus infection of rhesus macaques. J Virol. 2013;87(15):8351–62. [DOI] [PMC free article] [PubMed] [Google Scholar]
248.Wei C, Kakazu T, Chuah QY, Tanaka M, Kato G, Sano M. Reactivation of cyprinid herpesvirus 2 (CyHV-2) in asymptomatic surviving goldfish Carassius auratus (L.) under immunosuppression. Fish Shellfish Immunol. 2020;103:302–9. [DOI] [PubMed] [Google Scholar]
249.Thonur L, Russell GC, Stewart JP, Haig DM. Differential transcription of ovine herpesvirus 2 genes in lymphocytes from reservoir and susceptible species. Virus Genes. 2006;32(1):27–35. [DOI] [PubMed] [Google Scholar]
250.Perdrizet UG, Hill JE, Sobchishin L, Singh B, Fernando C, Bollinger TK, et al. Tissue and cellular tropism of Eptesicus fuscus gammaherpesvirus in big brown bats, potential role of pulmonary intravascular macrophages. Vet Pathol. 2024;61(4):550–61. [DOI] [PMC free article] [PubMed] [Google Scholar]
251.Mallet C, Cochard J, Leclercq S, Trapp-Fragnet L, Chouteau P, Denesvre C. Hypoxia and HIF-1 trigger Marek’s disease virus reactivation in lymphoma-derived latently infected T Lymphocytes. J Virol. 2022;96(5):e0142721. [DOI] [PMC free article] [PubMed] [Google Scholar]
252.Juillard F, Tan M, Li S, Kaye KM. Kaposi’s Sarcoma Herpesvirus Genome Persistence. Front Microbiol. 2016;7. Disponible sur: https://www.frontiersin.org/journals/microbiology/articles/10.3389/fmicb.2016.01149/full. Cité 24 févr 2025. [DOI] [PMC free article] [PubMed]
253.Kennedy PGE, Rovnak J, Badani H, Cohrs RJ. A comparison of herpes simplex virus type 1 and varicella-zoster virus latency and reactivation. J Gen Virol. 2015;96(Pt 7):1581–602. [DOI] [PMC free article] [PubMed] [Google Scholar]
254.Speck SH, Ganem D. Viral latency and its regulation: lessons from the γ-Herpesviruses. Cell Host Microbe. 2010;8(1):100–15. [DOI] [PMC free article] [PubMed] [Google Scholar]
255.Tempera I, Lieberman PM. Chromatin organization of gammaherpesvirus latent genomes. Biochim Biophys Acta. 2010;1799(3):236–45. [DOI] [PMC free article] [PubMed] [Google Scholar]
256.Tarrant-Elorza M, Rossetto CC, Pari GS. Maintenance and replication of the human cytomegalovirus genome during latency. Cell Host Microbe. 2014;16(1):43–54. [DOI] [PubMed] [Google Scholar]
257.Nair V. Latency and tumorigenesis in Marek’s disease. Avian Dis. 2013;57(2s1):360–5. [DOI] [PubMed] [Google Scholar]
258.Morissette G, Flamand L. Herpesviruses and chromosomal Integration. J Virol. 2010;84(23):12100–9. [DOI] [PMC free article] [PubMed] [Google Scholar]
259.Arbuckle JH, Medveczky MM, Luka J, Hadley SH, Luegmayr A, Ablashi D, et al. The latent human herpesvirus-6A genome specifically integrates in telomeres of human chromosomes in vivo and in vitro. Proc Natl Acad Sci. 2010;107(12):5563–8. [DOI] [PMC free article] [PubMed] [Google Scholar]
260.Kheimar A, Previdelli RL, Wight DJ, Kaufer BB. Telomeres and telomerase: role in marek’s disease virus pathogenesis, integration and tumorigenesis. Viruses. 2017;9(7):173. [DOI] [PMC free article] [PubMed] [Google Scholar]
261.Lestou VS, De Braekeleer M, Strehl S, Ott G, Gadner H, Ambros PF. Non-random integration of epstein-barr virus in lymphoblastoid cell lines. Genes Chromosom Cancer. 1993;8(1):38–48. [DOI] [PubMed] [Google Scholar]
262.Ahmed W, Liu ZF. Long Non-Coding RNAs: Novel Players in Regulation of Immune Response Upon Herpesvirus Infection. Frontiers in Immunology. 2018;9. Disponible sur: https://www.frontiersin.org/articles/10.3389/fimmu.2018.00761. Cité 19juill 2023. [DOI] [PMC free article] [PubMed]
263.Cliffe AR, Garber DA, Knipe DM. Transcription of the herpes simplex virus latency-associated transcript promotes the formation of facultative heterochromatin on lytic promoters. J Virol. 2009;83(16):8182–90. [DOI] [PMC free article] [PubMed] [Google Scholar]
264.Cullen BR. Viruses and microRNAs: RISCy interactions with serious consequences. Genes Dev. 2011;25(18):1881–94. [DOI] [PMC free article] [PubMed] [Google Scholar]
265.Pan D, Flores O, Umbach JL, Pesola JM, Bentley P, Rosato PC, et al. A neuron-specific host MicroRNA targets herpes simplex virus-1 ICP0 expression and promotes latency. Cell Host Microbe. 2014;15(4):446–56. [DOI] [PMC free article] [PubMed] [Google Scholar]
266.Grey F. Role of microRNAs in herpesvirus latency and persistence. J Gen Virol. 2015;96(4):739–51. [DOI] [PubMed] [Google Scholar]
267.Morgan RW, Burnside J. Roles of avian herpesvirus microRNAs in infection, latency, and oncogenesis. Biochim Biophys Acta. 2011;1809(11):654–9. [DOI] [PubMed] [Google Scholar]
268.Münz C. Latency and lytic replication in Epstein-Barr virus-associated oncogenesis. Nat Rev Microbiol. 2019;17(11):691–700. [DOI] [PubMed] [Google Scholar]
269.Chida Y, Mao X. Does psychosocial stress predict symptomatic herpes simplex virus recurrence? A meta-analytic investigation on prospective studies. Brain Behav Immun. 2009;23(7):917–25. [DOI] [PubMed] [Google Scholar]
270.El Hayderi L, Delvenne P, Rompen E, Senterre JM, Nikkels AF. Herpes simplex virus reactivation and dental procedures. Clin Oral Investig. 2013;17(8):1961–4. [DOI] [PubMed] [Google Scholar]
271.Padgett DA, Sheridan JF, Dorne J, Berntson GG, Candelora J, Glaser R. Social stress and the reactivation of latent herpes simplex virus type 1. Proc Natl Acad Sci. 1998;95(12):7231–5. [DOI] [PMC free article] [PubMed] [Google Scholar]
272.Roizman B, Whitley RJ. An inquiry into the molecular basis of HSV latency and reactivation. Annu Rev Microbiol. 2013;67:355–74. [DOI] [PubMed] [Google Scholar]
273.Suzich JB, Cliffe AR. Strength in diversity: Understanding the pathways to herpes simplex virus reactivation. Virology. 2018;522:81–91. [DOI] [PMC free article] [PubMed] [Google Scholar]
274.Tien YT, Akbar H, Jarosinski KW. Temperature-induced reactivation of Marek’s disease virus-transformed T cells ex vivo. Front Vet Sci. 2023;10:1145757. [DOI] [PMC free article] [PubMed] [Google Scholar]
275.Du T, Zhou G, Roizman B. HSV-1 gene expression from reactivated ganglia is disordered and concurrent with suppression of latency-associated transcript and miRNAs. Proc Natl Acad Sci. 2011;108(46):18820–4. [DOI] [PMC free article] [PubMed] [Google Scholar]
276.Kim JY, Mandarino A, Chao MV, Mohr I, Wilson AC. Transient reversal of episome silencing precedes VP16-dependent transcription during reactivation of latent HSV-1 in Neurons. PLoS Pathog. 2012;8(2):e1002540. [DOI] [PMC free article] [PubMed] [Google Scholar]
277.Hale AE, Moorman NJ. The ends dictate the means: promoter switching in herpesvirus gene expression. Annu Rev Virol. 2021;8:201–18. [DOI] [PubMed] [Google Scholar]
278.Jia Z, Zhang D, Zhu L, Xue J. Animal models of human herpesvirus infection. Anim Models Exp Med. 2025;8(4):615–28. [DOI] [PMC free article] [PubMed] [Google Scholar]
279.Petton B, Pernet F, Robert R, Boudry P. Temperature influence on pathogen transmission and subsequent mortalities in juvenile Pacific oysters Crassostrea gigas. Aquacult Environ Interact. 2013;3(3):257–73. [Google Scholar]
280.Rybak-Wolf A, Wyler E, Pentimalli TM, Legnini I, Oliveras Martinez A, Glažar P, et al. Modelling viral encephalitis caused by herpes simplex virus 1 infection in cerebral organoids. Nat Microbiol. 2023;8(7):1252–66. [DOI] [PMC free article] [PubMed] [Google Scholar]
281.Pourmaleki M, Jones CJ, Mellinghoff SD, Greenstein BD, Kumar P, Foronda M, et al. Multiplexed spatial profiling of hodgkin reed-sternberg cell neighborhoods in classic hodgkin lymphoma. Clin Cancer Res. 2024;30(17):3881–93. [DOI] [PMC free article] [PubMed] [Google Scholar]
282.Tombácz D, Balázs Z, Csabai Z, Moldován N, Szűcs A, Sharon D, et al. Characterization of the dynamic transcriptome of a herpesvirus with long-read single molecule real-time sequencing. Sci Rep. 2017;7(1):43751. [DOI] [PMC free article] [PubMed] [Google Scholar]
283.Rosani U, Bortoletto E, Zhang X, Huang BW, Xin LS, Krupovic M, et al. Long-read transcriptomics of Ostreid herpesvirus 1 uncovers a conserved expression strategy for the capsid maturation module and pinpoints a mechanism for evasion of the ADAR-based antiviral defence. Virus Evol. 2024;10(1):088. [DOI] [PMC free article] [PubMed] [Google Scholar]
284.Singh RK, Vangala R, Torne AS, Bose D, Robertson ES. Epigenetic and epitranscriptomic regulation during oncogenic γ-herpesvirus infection. Front Microbiol. 2024;15:1484455. [DOI] [PMC free article] [PubMed] [Google Scholar]
285.Sun A, Zhu X, Liu Y, Wang R, Yang S, Teng M, et al. Transcriptome-wide N6-methyladenosine modification profiling of long non-coding RNAs during replication of Marek’s disease virus in vitro. BMC Genomics. 2021;22(1):296. [DOI] [PMC free article] [PubMed] [Google Scholar]
286.Feng Z, Zhou F, Tan M, Wang T, Chen Y, Xu W, et al. Targeting m6A modification inhibits herpes virus 1 infection. Genes Dis. 2022;9(4):1114–28. [DOI] [PMC free article] [PubMed] [Google Scholar]
287.Gao C, Chen L, Tang SB, Long QY, He JL, Zhang NA, et al. The epigenetic landscapes of histone modifications on HSV-1 genome in human THP-1 cells. Antiviral Res. 2020;176:104730. [DOI] [PubMed] [Google Scholar]
288.Volkening JD, Spatz SJ, Ponnuraj N, Akbar H, Arrington JV, Vega-Rodriguez W, et al. Viral proteogenomic and expression profiling during productive replication of a skin-tropic herpesvirus in the natural host. PLoS Pathog. 2023;19(6):e1011204. [DOI] [PMC free article] [PubMed] [Google Scholar]
289.Jumper J, Evans R, Pritzel A, Green T, Figurnov M, Ronneberger O, et al. Highly accurate protein structure prediction with AlphaFold. Nature. 2021;596(7873):583–9. [DOI] [PMC free article] [PubMed] [Google Scholar]

Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Data Availability Statement

No datasets were generated or analysed during the current study.

[CR1] 1.Edwards RA, Rohwer F. Viral metagenomics. Nat Rev Microbiol juin. 2005;3(6):504–10. [DOI] [PubMed] [Google Scholar]

[CR2] 2.Hendrix RW, Smith MC, Burns RN, Ford ME, Hatfull GF. Evolutionary relationships among diverse bacteriophages and prophages: all the world’s a phage. Proc Natl Acad Sci U S A. 1999;96(5):2192–7. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR3] 3.Lawrence CM, Menon S, Eilers BJ, Bothner B, Khayat R, Douglas T, et al. Structural and functional studies of archaeal viruses. J Biol Chem. 2009;284(19):12599–603. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR4] 4.Nayfach S, Camargo AP, Schulz F, Eloe-Fadrosh E, Roux S, Kyrpides NC. CheckV assesses the quality and completeness of metagenome-assembled viral genomes | Nature Biotechnology. 2021. Disponible sur: https://www.nature.com/articles/s41587-020-00774-7. Cité 17 Nov 2024. [DOI] [PMC free article] [PubMed]

[CR5] 5.O’Leary NA, Wright MW, Brister JR, Ciufo S, Haddad D, McVeigh R, et al. Reference sequence (RefSeq) database at NCBI: current status, taxonomic expansion, and functional annotation. Nucleic Acids Res. 2016;44(D1):D733-745. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR6] 6.Silva JM, Pratas D, Caetano T, Matos S. The complexity landscape of viral genomes. GigaScience. 2022;11:giac079. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR7] 7.Gatherer D, Depledge DP, Hartley CA, Szpara ML, Vaz PK, Benkő M, et al. ICTV virus taxonomy profile: herpesviridae 2021. J Gen Virol. 2021;102(10):001673. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR8] 8.Knipe DM, Howley PM, Cohen JI, Griffin DE, Lamb RA, Martin MA, et al. Fields virology 6th edition. 2013.

[CR9] 9.Brito AF, Baele G, Nahata KD, Grubaugh ND, Pinney JW. Intrahost speciations and host switches played an important role in the evolution of herpesviruses. Virus Evol. 2021;7(1):veab025. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR10] 10.McGeoch DJ, Cook S, Dolan A, Jamieson FE, Telford EA. Molecular phylogeny and evolutionary timescale for the family of mammalian herpesviruses. J Mol Biol. 1995;247(3):443–58. [DOI] [PubMed] [Google Scholar]

[CR11] 11.McGeoch DJ, Dolan A, Ralph AC. Toward a comprehensive phylogeny for mammalian and avian herpesviruses. J Virol. 2000;74(22):10401–6. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR12] 12.McGeoch DJ, Cook S. Molecular phylogeny of the alphaherpesvirinae subfamily and a proposed evolutionary timescale. J Mol Biol. 1994;238(1):9–22. [DOI] [PubMed] [Google Scholar]

[CR13] 13.Davison AJ. Evolution of the herpesviruses. Vet Microbiol. 2002;86(1–2):69–88. [DOI] [PubMed] [Google Scholar]

[CR14] 14.Jackson A. The effect of paralogous lineages on the application of reconciliation analysis by cophylogeny mapping. Syst Biol. 2005;54(1):127–45. [DOI] [PubMed] [Google Scholar]

[CR15] 15.McGeoch DJ, Rixon FJ, Davison AJ. Topics in herpesvirus genomics and evolution. Virus Res. 2006;117(1):90–104. [DOI] [PubMed] [Google Scholar]

[CR16] 16.Escalera-Zamudio M, Rojas-Anaya E, Kolokotronis SO, Taboada B, Loza-Rubio E, Méndez-Ojeda ML, et al. Bats, primates, and the evolutionary origins and diversification of mammalian gammaherpesviruses. mBio. 2016;7(6):e01425. 10.1128/mbio.01425-16. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR17] 17.McGeoch DJ, Davison AJ, Dolan A, Gatherer D, Sevilla-Reyes EE. Molecular Evolution of the Herpesvirales. In: Domingo E, Parrish CR, Holland JJ, éditeurs. Origin and Evolution of Viruses (Second Edition). London: Academic Press; 2008. p. 447‑75. Disponible sur: https://www.sciencedirect.com/science/article/pii/B9780123741530000205. Cité 22 Janv 2025.

[CR18] 18.Fu M, Deng R, Wang J, Wang X. Whole-genome phylogenetic analysis of herpesviruses. Acta Virol. 2008;52(1):31–40. [PubMed] [Google Scholar]

[CR19] 19.Divilov K. Whole-genome assembly of a novel invertebrate herpesvirus from the gastropod Babylonia areolata. Microb Genom. 2024;10(4):001237. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR20] 20.Origgi FC, Schmidt BR, Lohmann P, Otten P, Meier RK, Pisano SRR, et al. Bufonid herpesvirus 1 (BfHV1) associated dermatitis and mortality in free ranging common toads (Bufo bufo) in Switzerland. Sci Rep. 2018;8(1):14737. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR21] 21.Walker L, Subramaniam K, Viadanna PHO, Vann JA, Marcquenski S, Godard D, et al. Characterization of an alloherpesvirus from wild lake sturgeon Acipenser fulvescens in Wisconsin (USA). Dis Aquat Organ. 2022;149:83–96. [DOI] [PubMed] [Google Scholar]

[CR22] 22.Henriquez RN, Polchana J, Kanchanakhan S, Davison AJ, Waltzek TB, Subramaniam K. Genome sequence of a novel alloherpesvirus isolated from glass catfish (Kryptopterus bicirrhis). Genome Announc. 2018;6(19):e00403. 10.1128/genomea.00403-18. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR23] 23.Davison AJ, Clements JB. Herpesviruses: General Properties. In: Topley & Wilson’s Microbiology and Microbial Infections. John Wiley & Sons, Ltd; 2010. Disponible sur: https://onlinelibrary.wiley.com/doi/abs/10.1002/9780470688618.taw0231. Cité 15 Jan 2025.

[CR24] 24.Cohen JI. Herpesvirus latency. American Society for Clinical Investigation; 2020. Disponible sur: https://www.jci.org/articles/view/136225/pdf. Cité 5 Jan 2023.

[CR25] 25.Everett RD. HSV-1 Biology and Life Cycle. In: Diefenbach RJ, Fraefel C, éditeurs. Herpes Simplex Virus: Methods and Protocols. New York, NY: Springer; 2014. p. 1‑17. (Methods in Molecular Biology). Disponible sur: 10.1007/978-1-4939-0428-0_1. Cité 17 juill 2023.

[CR26] 26.Griffin DE, Knipe D, Howley P, Lamb R, Martin M, Roizman B, et al. Fields virology. 2001.

[CR27] 27.Davison AJ. Herpesvirus systematics. Vet Microbiol. 2010;143(1):52–69. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR28] 28.Pellett PE, Roizman B. Chapter 59 : Herpesviridae. In: Fields Virology 6th edition. 2013.

[CR29] 29.Epstein MA. Observations on the mode of release of herpes virus from infected HeLa cells. J Cell Biol. 1962;12(3):589–97. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR30] 30.McCombs RM, Brunschwig P, Mirkovic R, Benyesh-Melnick M. Electron microscopic characterization of a herpeslike virus isolated from tree shrews. Virology. 1971;45(3):816–20. 10.1016/0042-6822(71)90203-0. [DOI] [PubMed]

[CR31] 31.Booy FP, Trus BL, Davison AJ, Steven AC. The capsid architecture of channel catfish virus, an evolutionarily distant herpesvirus, is largely conserved in the absence of discernible sequence homology with herpes simplex virus. Virology. 1996;215(2):134–41. [DOI] [PubMed] [Google Scholar]

[CR32] 32.Davison AJ, Trus BL, Cheng N, Steven AC, Watson MS, Cunningham C, et al. A novel class of herpesvirus with bivalve hosts. J Gen Virol. 2005;86(1):41–53. [DOI] [PubMed] [Google Scholar]

[CR33] 33.Liu F, Zhou ZH. Comparative virion structures of human herpesviruses. In: Arvin A, Campadelli-Fiume G, Mocarski E, Moore PS, Roizman B, Whitley R, et al., éditeurs. Human Herpesviruses: Biology, Therapy, and Immunoprophylaxis. Cambridge: Cambridge University Press; 2007. Disponible sur: http://www.ncbi.nlm.nih.gov/books/NBK47399/. Cité 18 Nov 2024.

[CR34] 34.Furlong D, Swift H, Roizman B. Arrangement of herpesvirus deoxyribonucleic acid in the core. J Virol. 1972;10(5):1071–4. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR35] 35.Nazerian K. DNA configuration in the core of Marek’s disease virus. J Virol mai. 1974;13(5):1148–50. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR36] 36.Zhou ZH, Chen DH, Jakana J, Rixon FJ, Chiu W. Visualization of tegument-capsid interactions and DNA in intact herpes simplex virus type 1 virions. J Virol. 1999;73(4):3210–8. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR37] 37.Heming JD, Conway JF, Homa FL. Herpesvirus Capsid Assembly and DNA Packaging. In: Osterrieder K, éditeur. Cell Biology of Herpes Viruses. Cham: Springer International Publishing; 2017. p. 119‑42. (Advances in Anatomy, Embryology and Cell Biology; vol. 223). Disponible sur: http://link.springer.com/10.1007/978-3-319-53168-7_6. Cité 20 avr 2023. [DOI] [PMC free article] [PubMed]

[CR38] 38.Johannsen E, Luftig M, Chase MR, Weicksel S, Cahir-McFarland E, Illanes D, et al. Proteins of purified Epstein-Barr virus. Proc Natl Acad Sci U S A. 2004;101(46):16286–91. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR39] 39.Kattenhorn LM, Mills R, Wagner M, Lomsadze A, Makeev V, Borodovsky M, et al. Identification of proteins associated with murine cytomegalovirus virions. J Virol. 2004;78(20):11187–97. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR40] 40.Michael K, Klupp BG, Mettenleiter TC, Karger A. Composition of pseudorabies virus particles lacking tegument protein US3, UL47, or UL49 or envelope glycoprotein E. J Virol. 2006;80(3):1332–9. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR41] 41.O’Connor CM, Kedes DH. Mass spectrometric analyses of purified rhesus monkey rhadinovirus reveal 33 virion-associated proteins. J Virol. 2006;80(3):1574–83. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR42] 42.Varnum SM, Streblow DN, Monroe ME, Smith P, Auberry KJ, Paša-Tolić L, et al. Identification of Proteins in Human Cytomegalovirus (HCMV) particles: the HCMV proteome. J Virol. 2004;78(20):10960–6. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR43] 43.Yuan S, Wang J, Zhu D, Wang N, Gao Q, Chen W, et al. Cryo-EM structure of a herpesvirus capsid at 3.1 Å. Science. 2018;360(6384):eaao7283. [DOI] [PubMed] [Google Scholar]

[CR44] 44.Wörner TP, Shamorkina TM, Snijder J, Heck AJR. Mass spectrometry-based structural virology. Anal Chem. 2021;93(1):620–40. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR45] 45.Eberle R, Black DH. Sequence of the ateline alphaherpesvirus 1 (HVA1) genome. Arch Virol. 2017;162(5):1423–5. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR46] 46.Pfaff F, Neubauer-Juric A, Krebs S, Hauser A, Singer S, Blum H, et al. Full genome sequence of bovine alphaherpesvirus 2 (BoHV-2). Arch Virol. 2021;166(2):639–43. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR47] 47.Tyler SD, Peters GA, Severini A. Complete genome sequence of cercopithecine herpesvirus 2 (SA8) and comparison with other simplexviruses. Virology. 2005;331:429. [DOI] [PubMed] [Google Scholar]

[CR48] 48.Davison AJ. Evolution of sexually transmitted and sexually transmissible human herpesviruses. Ann N Y Acad Sci. 2011;1230(1):E37-49. [DOI] [PubMed] [Google Scholar]

[CR49] 49.Babra B, Watson G, Xu W, Jeffrey BM, Xu JR, Rockey DD, et al. Analysis of the genome of leporid herpesvirus 4. Virology. 2012;433(1):183–91. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR50] 50.Perelygina L, Zhu L, Zurkuhlen H, Mills R, Borodovsky M, Hilliard JK. Complete sequence and comparative analysis of the genome of herpes B virus (Cercopithecine Herpesvirus 1) from a Rhesus Monkey. J Virol. 2003;77(11):6167–77. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR51] 51.Vaz PK, Mahony TJ, Hartley CA, Fowler EV, Ficorilli N, Lee SW, et al. The first genome sequence of a metatherian herpesvirus: macropodid herpesvirus 1. BMC Genomics. 2016;17(1):70. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR52] 52.Vaz PK, Mahony TJ, Hartley CA, Motha J, Devlin JM. Genome Sequences of Two Marsupial Simplex Viruses, Macropodid Alphaherpesviruses 2 and 4. Microbiol Resour Announc. 2021;10(2):e01189-20 Matthijnssens J, éditeur. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR53] 53.Severini A, Tyler SD, Peters GA, Black D, Eberle R. Genome sequence of a chimpanzee herpesvirus and its relation to other primate alphaherpesviruses. Arch Virol. 2013;158(8):1825–8. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR54] 54.Tyler SD, Severini A. The complete genome sequence of Herpesvirus Papio 2 (Cercopithecine Herpesvirus 16) shows evidence of recombination events among various progenitor herpesviruses. J Virol. 2006;80(3):1214–21. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR55] 55.Sasaki M, Setiyono A, Handharyani E, Kobayashi S, Rahmadani I, Taha S, et al. Isolation and characterization of a novel alphaherpesvirus in fruit bats. J Virol. 2014;88(17):9819–29. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR56] 56.Inagaki T, Yamada S, Fujii H, Yoshikawa T, Shibamura M, Harada S, et al. Characterization of a novel alphaherpesvirus isolated from the fruit bat pteropus lylei in Vietnam. J Virol. 2020;94(18):e00673-e720. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR57] 57.Tyler S, Severini A, Black D, Walker M, Eberle R. Structure and sequence of the saimiriine herpesvirus 1 genome. Virology. 2011;410(1):181–91. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR58] 58.Wang J, Höper D, Beer M, Osterrieder N. Complete genome sequence of virulent duck enteritis virus (DEV) strain 2085 and comparison with genome sequences of virulent and attenuated DEV strains. Virus Res. 2011;160(1):316–25. [DOI] [PubMed] [Google Scholar]

[CR59] 59.Guo Y, Li S, Sun X, He Y, Zhao H, Wang Y, et al. Complete genome sequence and evolution analysis of a columbid herpesvirus type 1 from feral pigeon in China. Arch Virol. 2017;162(7):2131–3. [DOI] [PubMed] [Google Scholar]

[CR60] 60.Tulman ER, Afonso CL, Lu Z, Zsak L, Rock DL, Kutish GF. The genome of a very virulent Marek’s disease virus. J Virol. 2000;74(17):7980–8. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR61] 61.Spatz SJ, Schat KA. Comparative genomic sequence analysis of the Marek’s disease vaccine strain SB-1. Virus Genes. 2011;42(3):331–8. [DOI] [PubMed] [Google Scholar]

[CR62] 62.Afonso CL, Tulman ER, Lu Z, Zsak L, Rock DL, Kutish GF. The genome of turkey herpesvirus. J Virol. 2001;75(2):971–8. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR63] 63.Pfaff F, Schulze C, König P, Franzke K, Bock S, Hlinak A, et al. A novel alphaherpesvirus associated with fatal diseases in banded Penguins. J Gen Virol. 2017;98(1):89–95. [DOI] [PubMed] [Google Scholar]

[CR64] 64.d’Offay JM, Fulton RW, Eberle R. Complete genome sequence of the NVSL BoHV-1.1 Cooper reference strain. Arch Virol. 2013;158(5):1109–13. [DOI] [PubMed] [Google Scholar]

[CR65] 65.Delhon G, Moraes MP, Lu Z, Afonso CL, Flores EF, Weiblen R, et al. Genome of bovine herpesvirus 5. J Virol. 2003;77(19):10339–47. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR66] 66.Scheffer CM, Varela APM, Cibulski SP, Schmidt C, Campos FS, Paim WP, et al. Genome sequence of bubaline alphaherpesvirus 1 (BuHV1) isolated in Australia in 1972. Arch Virol. 2017;162(5):1169–76. [DOI] [PubMed] [Google Scholar]

[CR67] 67.Papageorgiou KV, Suárez NM, Wilkie GS, McDonald M, Graham EM, Davison AJ. Genome sequence of canine herpesvirus. PLoS One. 2016;11(5):e0156015. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR68] 68.Hao F, Mao L, Li W, Li J, Yang L, Zhang W, et al. Epidemiological investigation and genomic characterization of Caprine herpesvirus 1 from goats in China. Infect Genet Evol. 2020;79:104168. [DOI] [PubMed] [Google Scholar]

[CR69] 69.Gray WL, Starnes B, White MW, Mahalingam R. The DNA sequence of the simian varicella virus genome. Virology. 2001;284(1):123–30. [DOI] [PubMed] [Google Scholar]

[CR70] 70.Telford EAR, Watson MS, McBride K, Davison AJ. The DNA sequence of equine herpesvirus-1. Virology. 1992;189(1):304–16. [DOI] [PubMed] [Google Scholar]

[CR71] 71.Sijmons S, Vissani A, Tordoya MS, Muylkens B, Thiry E, Maes P, et al. Complete genome sequence of equid Herpesvirus 3. Genome Announc. 2014;2(5):e00797-e814. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR72] 72.Telford EA, Watson MS, Perry J, Cullinane AA, Davison AJ. The DNA sequence of equine herpesvirus-4. J Gen Virol. 1998;79(5):1197–203. [DOI] [PubMed] [Google Scholar]

[CR73] 73.Martella V, Lanave G, Camero M, Larocca V, Lorusso E, Catella C, et al. Identification of a novel α-herpesvirus associated with ulcerative stomatitis in donkeys. Emerg Infect Dis. 2020;26(12):3044–7. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR74] 74.Garvey M, Suárez NM, Kerr K, Hector R, Moloney-Quinn L, Arkins S, et al. Equid herpesvirus 8: Complete genome sequence and association with abortion in mares. PLoS One. 2018;13(2):e0192301. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR75] 75.Kasem S, Yamada S, Kiupel M, Woodruff M, Ohya K, Fukushi H. Equine herpesvirus type 9 in giraffe with encephalitis. Emerg Infect Dis. 2008;14(12):1948–9. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR76] 76.Tai SHS, Niikura M, Cheng HH, Kruger JM, Wise AG, Maes RK. Complete genomic sequence and an infectious BAC clone of feline herpesvirus-1 (FHV-1). Virology. 2010;401(2):215–27. [DOI] [PubMed] [Google Scholar]

[CR77] 77.Davison AJ, Scott JE. The complete DNA sequence of varicella-zoster virus. J Gen Virol. 1986;67(Pt 9):1759–816. [DOI] [PubMed] [Google Scholar]

[CR78] 78.Davison AJ, Nielsen O, Subramaniam K, Jacob JM, Romero CH, Burek-Huntington KA, et al. Genome Sequence of an Alphaherpesvirus from a Beluga Whale (Delphinapterus leucas). Genome Announc. 2017;5(42):e01100-17. 10.1128/genomea.01100-17. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR79] 79.van Boheemen S, van Rijn AL, Pappas N, Carbo EC, Vorderman RHP, Sidorov I, et al. Retrospective validation of a metagenomic sequencing protocol for combined detection of RNA and DNA viruses using respiratory samples from pediatric patients. J Mol Diagn. 2020;22(2):196–207. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR80] 80.Szpara ML, Tafuri YR, Parsons L, Shamim SR, Verstrepen KJ, Legendre M, et al. A wide extent of inter-strain diversity in virulent and vaccine strains of alphaherpesviruses. PLoS Pathog. 2011;7(10):e1002282 Baines JD, éditeur. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR81] 81.Sutherland M, Sarker S, Vaz PK, Legione AR, Devlin JM, Macwhirter PL, et al. Disease surveillance in wild Victorian cacatuids reveals co-infection with multiple agents and detection of novel avian viruses. Vet Microbiol. 2019;235:257–64. [DOI] [PubMed] [Google Scholar]

[CR82] 82.Lee SW, Devlin JM, Markham JF, Noormohammadi AH, Browning GF, Ficorilli NP, et al. Comparative analysis of the complete genome sequences of two Australian origin live attenuated vaccines of infectious laryngotracheitis virus. Vaccine. 2011;29(52):9583–7. [DOI] [PubMed] [Google Scholar]

[CR83] 83.Thureen D, Keeler C. Psittacid Herpesvirus 1 and infectious laryngotracheitis virus: comparative genome sequence analysis of two avian alphaherpesviruses. J Virol. 2006;80:7863–72. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR84] 84.Sutherland M, Sarker S, Raidal SR. Molecular and microscopic characterisation of a novel pathogenic herpesvirus from Indian ringneck parrots (Psittacula krameri). Vet Microbiol. 2019;239:108428. [DOI] [PubMed] [Google Scholar]

[CR85] 85.Ackermann M, Koriabine M, Hartmann-Fritsch F, de Jong PJ, Lewis TD, Schetle N, et al. The genome of chelonid herpesvirus 5 harbors atypical genes. PLoS One. 2012;7(10):e46623. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR86] 86.Gandar F, Wilkie GS, Gatherer D, Kerr K, Marlier D, Diez M, et al. The Genome of a tortoise herpesvirus (Testudinid Herpesvirus 3) has a novel structure and contains a large region that is not required for replication in vitro or virulence in vivo. J Virol. 2015;89(22):11438–56. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR87] 87.Dolan A, Cunningham C, Hector RD, Hassan-Walker AF, Lee L, Addison C, et al. Genetic content of wild-type human cytomegalovirus. J Gen Virol. 2004;85(Pt 5):1301–12. [DOI] [PubMed] [Google Scholar]

[CR88] 88.Hansen SG, Strelow LI, Franchi DC, Anders DG, Wong SW. Complete sequence and genomic analysis of rhesus cytomegalovirus. J Virol. 2003;77(12):6620–36. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR89] 89.Marsh AK, Willer DO, Ambagala APN, Dzamba M, Chan JK, Pilon R, et al. Genomic sequencing and characterization of cynomolgus macaque cytomegalovirus. J Virol. 2011;85(24):12995–3009. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR90] 90.Blewett EL, Sherrod CJ, Texier JR, Conrad TM, Dittmer DP. Complete genome sequences of mandrillus leucophaeus and papio ursinus cytomegaloviruses. Genome Announc. 2015;3(4):e00781. 10.1128/genomea.00781-15. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR91] 91.Davison AJ, Dolan A, Akter P, Addison C, Dargan DJ, Alcendor DJ, et al. The human cytomegalovirus genome revisited: comparison with the chimpanzee cytomegalovirus genome. J Gen Virol. 2003;84(Pt 1):17–28. [DOI] [PubMed] [Google Scholar]

[CR92] 92.Cheng TP, Valentine MC, Gao J, Pingel JT, Yokoyama WM. Stability of murine cytomegalovirus genome after in vitro and in vivo passage. J Virol. 2010;84(5):2623–8. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR93] 93.Vink C, Beuken E, Bruggeman CA. Complete DNA sequence of the rat cytomegalovirus genome. J Virol. 2000;74(16):7656–65. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR94] 94.Ettinger J, Geyer H, Nitsche A, Zimmermann A, Brune W, Sandford GR, et al. Complete genome sequence of the english isolate of rat cytomegalovirus (Murid Herpesvirus 8). J Virol. 2012;86(24):13838–13838. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR95] 95.Yang D, Tamburro K, Dittmer D, Cui X, McVoy MA, Hernandez-Alvarado N, et al. Complete genome sequence of pathogenic guinea pig cytomegalovirus from salivary gland homogenates of infected animals. Genome Announc. 2013;1(2):e0005413. 10.1128/genomea.00054-13. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR96] 96.Zhang H, Todd S, Tachedjian M, Barr JA, Luo M, Yu M, et al. A novel bat herpesvirus encodes homologues of major histocompatibility complex classes I and II, C-type lectin, and a unique family of immune-related genes. J Virol. 2012;86(15):8014–30. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR97] 97.Bahr U, Darai G. Analysis and characterization of the complete genome of tupaia (tree shrew) herpesvirus. J Virol. 2001;75(10):4854–70. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR98] 98.Megaw AG, Rapaport D, Avidor B, Frenkel N, Davison AJ. The DNA sequence of the RK strain of human herpesvirus 7. Virology. 1998;244(1):119–32. [DOI] [PubMed] [Google Scholar]

[CR99] 99.Gompels UA, Nicholas J, Lawrence G, Jones M, Thomson BJ, Martin ME, et al. The DNA sequence of human herpesvirus-6: structure, coding content, and genome evolution. Virology. 1995;209(1):29–51. [DOI] [PubMed] [Google Scholar]

[CR100] 100.Dominguez G, Dambaugh TR, Stamey FR, Dewhurst S, Inoue N, Pellett PE. Human herpesvirus 6B genome sequence: coding content and comparison with human herpesvirus 6A. J Virol. 1999;73(10):8040–52. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR101] 101.Staheli JP, Dyen MR, Deutsch GH, Basom RS, Fitzgibbon MP, Lewis P, et al. Complete unique genome sequence, expression profile, and salivary gland tissue tropism of the herpesvirus 7 homolog in pigtailed macaques. J Virol. 2016;90(15):6657–74. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR102] 102.Patel SJ, Zhao G, Penna VR, Park E, Lauron EJ, Harvey IB, et al. A Murine Herpesvirus Closely Related to Ubiquitous Human Herpesviruses Causes T-Cell Depletion. J Virol. 2017;91(9):e02463-16 Sandri-Goldin RM, éditeur. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR103] 103.Wilkie GS, Davison AJ, Watson M, Kerr K, Sanderson S, Bouts T, et al. Complete genome sequences of elephant endotheliotropic herpesviruses 1A and 1B determined directly from fatal cases. J Virol. 2013;87(12):6700–12. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR104] 104.Long SY, Latimer EM, Hayward GS. Review of elephant endotheliotropic herpesviruses and acute hemorrhagic disease. ILAR J. 2016;56(3):283–96. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR105] 105.Ling PD, Long SY, Fuery A, Peng RS, Heaggans SY, Qin X, et al. Complete Genome Sequence of Elephant Endotheliotropic Herpesvirus 4, the First Example of a GC-Rich Branch Proboscivirus. mSphere. 2016;1(3):e00081-15. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR106] 106.Davison AJ, Subramaniam K, Kerr K, Jacob JM, Landrau-Giovannetti N, Walsh MT, et al. Genome Sequence of a Gammaherpesvirus from a Common Bottlenose Dolphin (Tursiops truncatus). Genome Announc. 2017;5(31):e00777-e817. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR107] 107.Ensser A, Pflanz R, Fleckenstein B. Primary structure of the alcelaphine herpesvirus 1 genome. J Virol. 1997;71(9):6517–25. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR108] 108.Taus NS, O’Toole D, Herndon DR, Cunha CW, Warg JV, Seal BS, et al. Malignant catarrhal fever in American bison (Bison bison) experimentally infected with alcelaphine herpesvirus 2. Vet Microbiol. 2014;172(1):318–22. [DOI] [PubMed] [Google Scholar]

[CR109] 109.Jia J, Delhon G, Tulman ER, Diel DG, Osorio FA, Wen X, et al. Novel gammaherpesvirus functions encoded by bovine herpesvirus 6 (bovine lymphotropic virus). J Gen Virol. 2014;95(8):1790–8. [DOI] [PubMed] [Google Scholar]

[CR110] 110.Chmielewicz B, Goltz M, Ehlers B. Detection and multigenic characterization of a novel gammaherpesvirus in goats. Virus Res. 2001;75(1):87–94. [DOI] [PubMed] [Google Scholar]

[CR111] 111.Li H, Gailbreath K, Flach EJ, Taus NS, Cooley J, Keller J, et al. A novel subgroup of rhadinoviruses in ruminants. J Gen Virol. 2005;86(Pt 11):3021–6. [DOI] [PubMed] [Google Scholar]

[CR112] 112.Hart J, Ackermann M, Jayawardane G, Russell G, Haig DM, Reid H, et al. Complete sequence and analysis of the ovine herpesvirus 2 genome. J Gen Virol. 2007;88(1):28–39. [DOI] [PubMed] [Google Scholar]

[CR113] 113.Ehlers B, Ulrich S, Goltz M. Detection of two novel porcine herpesviruses with high similarity to gammaherpesviruses. J Gen Virol. 1999;80(4):971–8. [DOI] [PubMed] [Google Scholar]

[CR114] 114.Chmielewicz B, Goltz M, Franz T, Bauer C, Brema S, Ellerbrok H, et al. A novel porcine gammaherpesvirus. Virology. 2003;308(2):317–29. [DOI] [PubMed] [Google Scholar]

[CR115] 115.Telford EAR, Watson MS, Aird HC, Perry J, Davison AJ. The DNA sequence of equine herpesvirus 2. J Mol Biol. 1995;249(3):520–8. [DOI] [PubMed] [Google Scholar]

[CR116] 116.Wilkie GS, Kerr K, Stewart JP, Studdert MJ, Davison AJ. Genome Sequences of equid herpesviruses 2 and 5. Genome Announce. 2015;3(2):e00119. 10.1128/genomea.00119-15. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR117] 117.Troyer RM, Lee JS, Vuyisich M, Chain P, Lo CC, Kronmiller B, et al. First complete genome sequence of felis catus gammaherpesvirus 1. Genome Announc. 2015;3(6):e01192. 10.1128/genomea.01192-15. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR118] 118.Banks M, King DP, Daniells C, Stagg DA, Gavier-Widen D. Partial characterization of a novel gammaherpesvirus isolated from a European badger (Meles meles). J Gen Virol. 2002;83(Pt 6):1325–30. [DOI] [PubMed] [Google Scholar]

[CR119] 119.Bellehumeur C, Nielsen O, Measures L, Harwood L, Goldstein T, Boyle B, et al. Herpesviruses including novel gammaherpesviruses are widespread among phocid seal species in Canada. J Wildl Dis. 2016;52(1):70–81. [DOI] [PubMed] [Google Scholar]

[CR120] 120.Noguchi K, Kuwata R, Shimoda H, Mizutani T, Hondo E, Maeda K. The complete genomic sequence of Rhinolophus gammaherpesvirus 1 isolated from a greater horseshoe bat. Arch Virol. 2019;164(1):317–9. [DOI] [PubMed] [Google Scholar]

[CR121] 121.Shabman RS, Shrivastava S, Tsibane T, Attie O, Jayaprakash A, Mire CE, et al. Isolation and characterization of a novel gammaherpesvirus from a microbat cell line. mSphere. 2016;1(1):e00070-15. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR122] 122.Richter J, Puchtler I, Fleckenstein B. Thymidylate synthase gene of herpesvirus ateles. J Virol. 1988;62(9):3530–5. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR123] 123.Albrecht JC. Primary structure of the Herpesvirus ateles genome. J Virol. 2000;74(2):1033–7. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR124] 124.Zimmermann W, Broll H, Ehlers B, Buhk HJ, Rosenthal A, Goltz M. Genome sequence of bovine herpesvirus 4, a bovine Rhadinovirus, and identification of an origin of DNA replication. J Virol. 2001;75(3):1186–94. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR125] 125.Dhingra A, Ganzenmueller T, Hage E, Suárez NM, Mätz-Rensing K, Widmer D, et al. Novel Virus Related to Kaposi’s Sarcoma–Associated Herpesvirus from Colobus Monkey - Volume 25, Number 8—August 2019 - Emerging Infectious Diseases journal - CDC. 2019; Disponible sur: https://wwwnc.cdc.gov/eid/article/25/8/18-1802_article. Cité 16 Juill 2024. [DOI] [PMC free article] [PubMed]

[CR126] 126.Loh J, Zhao G, Nelson CA, Coder P, Droit L, Handley SA, et al. Identification and sequencing of a novel rodent gammaherpesvirus that establishes acute and latent infection in laboratory mice. J Virol. 2011;85(6):2642–56. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR127] 127.Glenn M, Rainbow L, Auradé F, Davison A, Schulz TF. Identification of a spliced gene from Kaposi’s sarcoma-associated herpesvirus encoding a protein with similarities to latent membrane proteins 1 and 2A of Epstein-Barr virus. J Virol. 1999;73(8):6953–63. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR128] 128.Axthelm MK, Bourdette DN, Marracci GH, Su W, Mullaney ET, Manoharan M, et al. Japanese macaque encephalomyelitis: a spontaneous multiple sclerosis–like disease in a nonhuman primate. Ann Neurol. 2011;70(3):362–73. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR129] 129.Bruce AG, Thouless ME, Haines AS, Pallen MJ, Grundhoff A, Rose TM. Complete genome sequence of Pig-tailed macaque rhadinovirus 2 and its evolutionary relationship with rhesus macaque rhadinovirus and human herpesvirus 8/Kaposi’s sarcoma-associated herpesvirus. J Virol. 2015;89(7):3888–909. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR130] 130.Searles RP, Bergquam EP, Axthelm MK, Wong SW. Sequence and Genomic Analysis of a Rhesus Macaque Rhadinovirus with Similarity to Kaposi’s Sarcoma-Associated Herpesvirus/Human Herpesvirus 8. J Virol. 1999;73(4):3040–53. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR131] 131.Bruce AG, Ryan JT, Thomas MJ, Peng X, Grundhoff A, Tsai CC, et al. Next-generation sequence analysis of the genome of RFHVMn, the macaque homolog of Kaposi’s sarcoma (KS)-associated herpesvirus, from a KS-like tumor of a pig-tailed macaque. J Virol. 2013;87(24):13676–93. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR132] 132.Virgin HW, Latreille P, Wamsley P, Hallsworth K, Weck KE, Dal Canto AJ, et al. Complete sequence and genomic analysis of murine gammaherpesvirus 68. J Virol. 1997;71(8):5894–904. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR133] 133.Hughes DJ, Kipar A, Milligan SG, Cunningham C, Sanders M, Quail MA, et al. Characterization of a novel wood mouse virus related to murid herpesvirus 4. J Gen Virol. 2010;91(Pt 4):867–79. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR134] 134.Albrecht JC, Nicholas J, Biller D, Cameron KR, Biesinger B, Newman C, et al. Primary structure of the herpesvirus saimiri genome. J Virol. 1992;66(8):5047–58. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR135] 135.Subudhi S, Rapin N, Dorville N, Hill JE, Town J, Willis CKR, et al. Isolation, characterization and prevalence of a novel Gammaherpesvirus in Eptesicus fuscus, the North American big brown bat. Virology. 2018;516:227–38. [DOI] [PubMed] [Google Scholar]

[CR136] 136.Vaz PK, Hartley CA, Lee SY, Sansom FM, Adams TE, Stalder K, et al. Koala and Wombat Gammaherpesviruses Encode the First Known Viral NTPDase Homologs and Are Phylogenetically Divergent from All Known Gammaherpesviruses. J Virol. 2019;93(6):e01404-e1418. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR137] 137.Cho Y, Ramer J, Rivailler P, Quink C, Garber RL, Beier DR, et al. An Epstein-Barr-related herpesvirus from marmoset lymphomas. Proc Natl Acad Sci U S A. 2001;98(3):1224–9. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR138] 138.Gerner CS, Dolan A, McGeoch DJ. Phylogenetic relationships in the Lymphocryptovirus genus of the Gammaherpesvirinae. Virus Res. 2004;99(2):187–92. [DOI] [PubMed] [Google Scholar]

[CR139] 139.Baer R, Bankier AT, Biggin MD, Deininger PL, Farrell PJ, Gibson TJ, et al. DNA sequence and expression of the B95–8 Epstein—Barr virus genome. Nature. 1984;310(5974):207–11. [DOI] [PubMed] [Google Scholar]

[CR140] 140.Kamperschroer C, Gosink MM, Kumpf SW, O’Donnell LM, Tartaro KR. The genomic sequence of lymphocryptovirus from cynomolgus macaque. Virology. 2016;488:28–36. [DOI] [PubMed] [Google Scholar]

[CR141] 141.Krumbholz A, Roempke J, Liehr T, Groth M, Meerbach A, Schacke M, et al. Macaca arctoides gammaherpesvirus 1 (strain herpesvirus Macaca arctoides): virus sequence, phylogeny and characterisation of virus-transformed macaque and rabbit cell lines. Med Microbiol Immunol. 2019;208(1):109–29. [DOI] [PubMed] [Google Scholar]

[CR142] 142.Rivailler P, Jiang H, Cho YG, Quink C, Wang F. Complete Nucleotide Sequence of the Rhesus Lymphocryptovirus: Genetic Validation for an Epstein-Barr Virus Animal Model. J Virol. 2002;76(1):421–6. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR143] 143.Savin KW, Cocks BG, Wong F, Sawbridge T, Cogan N, Savage D, et al. A neurotropic herpesvirus infecting the gastropod, abalone, shares ancestry with oyster herpesvirus and a herpesvirus associated with the amphioxus genome. Virol J. 2010;7(1):308. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR144] 144.Davison AJ, Cunningham C, Sauerbier W, McKinnell RG. Genome sequences of two frog herpesviruses. J Gen Virol. 2006;87(12):3509–14. [DOI] [PubMed] [Google Scholar]

[CR145] 145.Origgi FC, Schmidt BR, Lohmann P, Otten P, Akdesir E, Gaschen V, et al. Ranid herpesvirus 3 and proliferative dermatitis in free-ranging wild common frogs (Rana Temporaria). Vet Pathol. 2017;54(4):686–94. [DOI] [PubMed] [Google Scholar]

[CR146] 146.Doszpoly A, Kovács ER, Bovo G, LaPatra SE, Harrach B, Benko M. Molecular confirmation of a new herpesvirus from catfish (Ameiurus melas) by testing the performance of a novel PCR method, designed to target the DNA polymerase gene of alloherpesviruses. Arch Virol. 2008;153(11):2123–7. [DOI] [PubMed] [Google Scholar]

[CR147] 147.Davison AJ. Channel catfish virus: a new type of herpesvirus. Virology. 1992;186(1):9–14. [DOI] [PubMed] [Google Scholar]

[CR148] 148.Borzák R, Haluk T, Bartha D, Doszpoly A. Complete genome sequence and analysis of ictalurid herpesvirus 2. Arch Virol. 2018;163(4):1083–5. [DOI] [PubMed] [Google Scholar]

[CR149] 149.van Beurden SJ, Bossers A, Voorbergen-Laarman MHA, Haenen OLM, Peters S, Abma-Henkens MHC, et al. Complete genome sequence and taxonomic position of anguillid herpesvirus 1. J Gen Virol. 2010;91(4):880–7. [DOI] [PubMed] [Google Scholar]

[CR150] 150.Davison AJ, Kurobe T, Gatherer D, Cunningham C, Korf I, Fukuda H, et al. Comparative genomics of carp herpesviruses. J Virol. 2013;87(5):2908–22. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR151] 151.Aoki T, Hirono I, Kurokawa K, Fukuda H, Nahary R, Eldar A, et al. Genome sequences of three koi herpesvirus isolates representing the expanding distribution of an emerging disease threatening koi and common carp worldwide. J Virol. 2007;81(10):5058–65. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR152] 152.Davison AJ. The genome of salmonid herpesvirus 1. J Virol. 1998;72(3):1974–82. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR153] 153.Waltzek T, Kelley G, Alfaro M, Kurobe T, Davison A, Hedrick R. Phylogenetic relationships in the family Alloherpesviridae. Dis Aquat Org. 2009;84:179–94. [DOI] [PubMed] [Google Scholar]

[CR154] 154.Boutier M, Morvan L, Delrez N, Origgi F, Doszpoly A, Vanderplasschen A. Fish and Amphibian Alloherpesviruses (Herpesviridae). In: Encyclopedia of Virology. Elsevier; 2021. p. 306‑15. Disponible sur: https://linkinghub.elsevier.com/retrieve/pii/B978012809633820931X. Cité 8 août 2023.

[CR155] 155.Rosani U, Gaia M, Delmont TO, Krupovic M. Tracing the invertebrate herpesviruses in the global sequence datasets. Front Mar Sci. 2023;10:1159754. [Google Scholar]

[CR156] 156.Ferreira RC, Alves GV, Ramon M, Antoneli F, Briones MRS. Reconstructing prehistoric viral genomes from neanderthal sequencing data. Viruses. 2024;16(6):856. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR157] 157.Alfano N, Dayaram A, Axtner J, Tsangaras K, Kampmann ML, Mohamed A, et al. Non-invasive surveys of mammalian viruses using environmental DNA. Methods Ecol Evol. 2021;12(10):1941–52. [Google Scholar]

[CR158] 158.Bass D, Christison KW, Stentiford GD, Cook LSJ, Hartikainen H. Environmental DNA/RNA for pathogen and parasite detection, surveillance, and ecology. Trends Parasitol. 2023;39(4):285–304. [DOI] [PubMed] [Google Scholar]

[CR159] 159.Bavarsad M, Abed-Elmdoost A, Tabandeh MR, Alishahi M, Mirvaghefi A, Farahmand H. Rapid cyprinid herpesvirus 3 detection using loop-mediated isothermal amplification (LAMP) combined with gold nanoprobes and SYBR safe. Aquaculture. 2025;594:741374. [Google Scholar]

[CR160] 160.Dotto-Maurel A, Pelletier C, Morga B, Jacquot M, Faury N, Dégremont L, et al. Evaluation of tangential flow filtration coupled to long-read sequencing for ostreid herpesvirus type 1 genome assembly. Microbial Genomics. 2022;8(11). Disponible sur: https://www.microbiologyresearch.org/content/journal/mgen/. 10.1099/mgen.0.000895. Cité 14 Nov 2022. [DOI] [PMC free article] [PubMed]

[CR161] 161.Hanson L, Dishon A, Kotler M. Herpesviruses that infect fish. Viruses. 2011;3(11):2160–91. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR162] 162.Khanthaphixay B, Wu L, Yoon JY. Microparticle-based detection of viruses. Biosensors. 2023;13(8):820. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR163] 163.Volpe E, Errani F, Mandrioli L, Ciulli S. Advances in viral aquatic animal disease knowledge: the molecular methods’ contribution. Biology mars. 2023;12(3):466. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR164] 164.Gu W, Zeng N, Zhou L, Ge X, Guo X, Yang H. Genomic organization and molecular characterization of porcine cytomegalovirus. Virology. 2014;460–461:165–72. [DOI] [PubMed] [Google Scholar]

[CR165] 165.Chang W, Jiao X, Sui H, Goswami S, Sherman BT, Fromont C, et al. Complete genome sequence of herpes simplex virus 2 strain G. Viruses. 2022;14(3):536. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR166] 166.Delius H, Clements JB. A partial denaturation map of herpes simplex virus Type 1 DNA: evidence for inversions of the unique DNA Regions. J Gen Virol. 1976;33(1):125–33. [DOI] [PubMed] [Google Scholar]

[CR167] 167.Desrosiers RC, Falk LA. Herpesvirus tamarinus and its relation to herpes simplex virus. J Gen Virol. 1981;56(1):119–30. [DOI] [PubMed] [Google Scholar]

[CR168] 168.Henry BE, Robinson RA, Dauenhauer SA, Atherton SS, Hayward GS, O’Callaghan DJ. Structure of the genome of equine herpesvirus type 1. Virology. 1981;115(1):97–114. [DOI] [PubMed] [Google Scholar]

[CR169] 169.Jenkins FJ, Roizman B. Herpes simplex virus 1 recombinants with noninverting genomes frozen in different isomeric arrangements are capable of independent replication. J Virol. 1986;59(2):494–9. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR170] 170.Leib DA, Bradbury JM, Hart CA, McCarthy K. Genome isomerism in two alphaherpesviruses: Herpesvirus saimiri-1(Herpesvirus tamarinus) and avian infectious laryngotracheitis virus. Adv Virol. 1987;93(3):287–94. [DOI] [PubMed] [Google Scholar]

[CR171] 171.Lomniczi B, Blankenship ML, Ben-Porat T. Deletions in the genomes of pseudorabies virus vaccine strains and existence of four isomers of the genomes. J Virol. 1984;49(3):970–9. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR172] 172.Sauer A, Wang JB, Hahn G, McVoy MA. A human cytomegalovirus deleted of internal repeats replicates with near wild type efficiency but fails to undergo genome isomerization. Virology. 2010;401(1):90–5. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR173] 173.Schynts F, McVoy MA, Meurens F, Detry B, Epstein AL, Thiry E. The structures of bovine herpesvirus 1 virion and concatemeric DNA: implications for cleavage and packaging of herpesvirus genomes. Virology. 2003;314(1):326–35. [DOI] [PubMed] [Google Scholar]

[CR174] 174.Wood ML, Neumann R, Roy P, Nair V, Royle NJ. Characterization of integrated Marek’s disease virus genomes supports a model of integration by homology-directed recombination and telomere-loop-driven excision. J Virol. 2023;97(10):e00716-e723. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR175] 175.Mahiet C, Ergani A, Huot N, Alende N, Azough A, Salvaire F, et al. Structural variability of the herpes simplex virus 1 genome in vitro and in vivo. J Virol. 2012;86(16):8592–601. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR176] 176.Kinchington PR, Reinhold WC, Casey TA, Straus SE, Hay J, Ruyechan WT. Inversion and circularization of the varicella-zoster virus genome. J Virol. 1985;56(1):194–200. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR177] 177.Nicolson L. The Characterisation of Specific EHV-4 Genes [PhD]. [Ann Arbor :]: ProQuest Dissertations & Theses; 1991. Disponible sur: https://theses.gla.ac.uk/77027/. Cité 11 Sept 2023.

[CR178] 178.Weber PC, Challberg MD, Nelson NJ, Levine M, Glorioso JC. Inversion events in the HSV-1 genome are directly mediated by the viral DNA replication machinery and lack sequence specificity. Cell. 1988;54(3):369–81. [DOI] [PubMed] [Google Scholar]

[CR179] 179.Jacob RJ, Morse LS, Roizman B. Anatomy of herpes simplex virus DNA XII. Accumulation of head-to-tail concatemers in nuclei of infected cells and their role in the generation of the four isomeric arrangements of viral DNA. J Virol. 1979;29(2):448–57. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR180] 180.Roizman B. The structure and isomerization of herpes simplex virus genomes. Cell. 1979;16(3):481–94. [DOI] [PubMed] [Google Scholar]

[CR181] 181.Hayward GS, Jacob RJ, Wadsworth SC, Roizman B. Anatomy of herpes simplex virus DNA: evidence for four populations of molecules that differ in the relative orientations of their long and short components. 1975. Disponible sur: https://www.pnas.org/doi/10.1073/pnas.72.11.4243. Cité 3 Sept 2023. [DOI] [PMC free article] [PubMed]

[CR182] 182.McKenzie J, El-Guindy A. Epstein-Barr Virus Lytic Cycle Reactivation. In: Münz C, éditeur. Epstein Barr Virus Volume 2: One Herpes Virus: Many Diseases. Cham: Springer International Publishing; 2015. p. 237‑61. (Current Topics in Microbiology and Immunology). Disponible sur: 10.1007/978-3-319-22834-1_8. Cité 17 juill 2023. [DOI] [PubMed]

[CR183] 183.Boldogh I, Albrecht T, Porter DD. Persistent Viral Infections. In: Baron S, éditeur. Medical Microbiology. 4th éd. Galveston (TX): University of Texas Medical Branch at Galveston; 1996. Disponible sur: http://www.ncbi.nlm.nih.gov/books/NBK8538/. Cité 27 Juill 2023. [PubMed]

[CR184] 184.Brunner S, Herndler-Brandstetter D, Weinberger B, Grubeck-Loebenstein B. Persistent viral infections and immune aging. Ageing Res Rev. 2011;10(3):362–9. [DOI] [PubMed] [Google Scholar]

[CR185] 185.Frebel H, Richter K, Oxenius A. How chronic viral infections impact on antigen-specific T-cell responses. Eur J Immunol. 2010;40(3):654–63. [DOI] [PubMed] [Google Scholar]

[CR186] 186.Gerna G, Kabanova A, Lilleri D. Human cytomegalovirus cell tropism and host cell receptors. Vaccines (Basel). 2019;7(3):70. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR187] 187.Wofford AS, McCusker I, Green JC, Vensko TA, Pellett PE. Chapter Ten - Betaherpesvirus assembly and egress: Recent advances illuminate the path. In: Kielian M, Mettenleiter TC, Roossinck MJ, éditeurs. Advances in Virus Research. Academic Press; 2020. p. 337‑92. (Virus Assembly and Exit Pathways; vol. 108). Disponible sur: https://www.sciencedirect.com/science/article/pii/S0065352720300439. Cité 25 Nov 2024. [DOI] [PubMed]

[CR188] 188.Liashkovich I, Hafezi W, Kühn JM, Oberleithner H, Shahin V. Nuclear delivery mechanism of herpes simplex virus type 1 genome. J Mol Recognit. 2011;24(3):414–21. [DOI] [PubMed] [Google Scholar]

[CR189] 189.Kukhanova MK, Korovina AN, Kochetkov SN. Human herpes simplex virus: Life cycle and development of inhibitors. Biochemistry Moscow. 2014;79(13):1635–52. [DOI] [PubMed] [Google Scholar]

[CR190] 190.Castillo JP, Kowalik TF. HCMV infection: modulating the cell cycle and cell death. Int Rev Immunol. 2004;23(1–2):113–39. [DOI] [PubMed] [Google Scholar]

[CR191] 191.Damania B, Kenney SC, Raab-Traub N. Epstein-Barr virus: biology and clinical disease. Cell. 2022;185(20):3652–70. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR192] 192.Boehmer P, Nimonkar A. Herpes virus replication. IUBMB Life. 2003;55(1):13–22. [DOI] [PubMed] [Google Scholar]

[CR193] 193.Zhu S, Viejo-Borbolla A. Pathogenesis and virulence of herpes simplex virus. Virulence. 2021;12(1):2670–702. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR194] 194.DeLuca NA, Schaffer PA. Activation of immediate-early, early, and late promoters by temperature-sensitive and wild-type forms of herpes simplex virus type 1 protein ICP4. Mol Cell Biol. 1985;5(8):1997–2008. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR195] 195.Hardwicke MA, Sandri-Goldin RM. The herpes simplex virus regulatory protein ICP27 contributes to the decrease in cellular mRNA levels during infection. J Virol. 1994;68(8):4797–810. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR196] 196.Spencer CA, Dahmus ME, Rice SA. Repression of host RNA polymerase II transcription by herpes simplex virus type 1. J Virol. 1997;71(3):2031–40. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR197] 197.Homa FL, Brown JC. Capsid assembly and DNA packaging in herpes simplex virus. Rev Med Virol. 1997;7(2):107–22. [DOI] [PubMed] [Google Scholar]

[CR198] 198.Close WL, Anderson AN, Pellett PE. Betaherpesvirus Virion Assembly and Egress. In: Kawaguchi Y, Mori Y, Kimura H, éditeurs. Human Herpesviruses. Singapore: Springer; 2018. p. 167‑207. Disponible sur: 10.1007/978-981-10-7230-7_9. Cité 17 Nov 2024.

[CR199] 199.Crump C. Virus assembly and egress of HSV. Adv Exp Med Biol. 2018;1045:23–44. [DOI] [PubMed] [Google Scholar]

[CR200] 200.Hellberg T, Paßvogel L, Schulz KS, Klupp BG, Mettenleiter TC. Nuclear egress of herpesviruses: the prototypic vesicular nucleocytoplasmic transport. Adv Virus Res. 2016;94:81–140. [DOI] [PubMed] [Google Scholar]

[CR201] 201.Roller RJ, Baines JD. Herpesvirus nuclear egress. Adv Anat Embryol Cell Biol. 2017;223:143–69. [DOI] [PubMed] [Google Scholar]

[CR202] 202.Dodding MP, Way M. Coupling viruses to dynein and kinesin‐1. The EMBO Journal. 2011; Disponible sur: https://www.embopress.org/doi/10.1038/emboj.2011.283. Cité 25 2024. [DOI] [PMC free article] [PubMed]

[CR203] 203.Lyman MG, Enquist LW. Herpesvirus interactions with the host cytoskeleton. J Virol. 2009;83(5):2058–66. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR204] 204.Naghavi MH, Walsh D. Microtubule regulation and function during virus infection. J Virol. 2017;91(16):e00538-e617. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR205] 205.Abdelgawad A, Damiani A, Ho SYW, Strauss G, Szentiks CA, East ML, et al. Zebra Alphaherpesviruses (EHV-1 and EHV-9): genetic diversity, latency and co-infections. Viruses. 2016;8(9):262. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR206] 206.Aghazadeh M, Shi M, Pesavento PA, Durham AC, Polley T, Donahoe SL, et al. Transcriptome analysis and in situ hybridization for FcaGHV1 in feline lymphoma. Viruses. 2018;10(9):464. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR207] 207.Aligo J, Walker M, Bugelski P, Weinstock D. Is murine gammaherpesvirus-68 (MHV-68) a suitable immunotoxicological model for examining immunomodulatory drug-associated viral recrudescence? J Immunotoxicol. 2015;12(1):1–15. [DOI] [PubMed] [Google Scholar]

[CR208] 208.Bielefeldt-Ohmann H, Bruce AG, Howard K, Ikoma M, Thouless ME, Rose TM. Macaque homologs of Kaposi’s sarcoma-associated herpesvirus (KSHV) infect germinal center lymphoid cells, epithelial cells in skin and gastrointestinal tract and gonadal germ cells in naturally infected macaques. Virology. 2018;519:106–20. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR209] 209.Broussard G, Damania B. Regulation of KSHV latency and lytic reactivation. Viruses. 2020;12(9):1034. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR210] 210.Camero M, Lanave G, Lucente MS, Losurdo M, Di Paola G, Lorusso E, et al. Bubaline alphaherpesvirus 1 induces a latent/reactivable infection in goats. Comp Immunol Microbiol Infect Dis. 2019;62:54–7. [DOI] [PubMed] [Google Scholar]

[CR211] 211.Campos FS, Franco AC, Oliveira MT, Firpo R, Strelczuk G, Fontoura FE, et al. Detection of bovine herpesvirus 2 and bovine herpesvirus 4 DNA in trigeminal ganglia of naturally infected cattle by polymerase chain reaction. Vet Microbiol. 2014;171(1):182–8. [DOI] [PubMed] [Google Scholar]

[CR212] 212.Collins CM, Medveczky MM, Lund T, Medveczky PG. The terminal repeats and latency-associated nuclear antigen of herpesvirus saimiri are essential for episomal persistence of the viral genome. J Gen Virol. 2002;83(9):2269–78. [DOI] [PubMed] [Google Scholar]

[CR213] 213.Darai G, Koch HG. Tree Shrew Herpesvirus: Pathogenicity and Latency. In: Wittmann G, Gaskell RM, Rziha HJ, éditeurs. Latent Herpes Virus Infections in Veterinary Medicine: A Seminar in the CEC Programme of Coordination of Research on Animal Pathology, held at Tübingen, Federal Republic of Germany, September 21–24, 1982. Dordrecht: Springer Netherlands; 1984. p. 91‑102. Disponible sur: 10.1007/978-94-009-5662-9_8. Cité 19 févr 2025.

[CR214] 214.das Neves CG, Mørk T, Thiry J, Godfroid J, Rimstad E, Thiry E, et al. Cervid herpesvirus 2 experimentally reactivated in reindeer can produce generalized viremia and abortion. Virus Res. 2009;145(2):321–8. [DOI] [PubMed] [Google Scholar]

[CR215] 215.Denesvre C, You Y, Rémy S, Vychodil T, Courvoisier K, Penzes Z, et al. Impact of viral telomeric repeat sequences on herpesvirus vector vaccine integration and persistence. PLOS Pathogens. 2024;20(5):e1012261. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR216] 216.Dewals BG, Vanderplasschen A. Malignant catarrhal fever induced by Alcelaphine herpesvirus 1 is characterized by an expansion of activated CD3+CD8+CD4- T cells expressing a cytotoxic phenotype in both lymphoid and non-lymphoid tissues. Vet Res. 2011;42(1):95. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR217] 217.Divilov K, Wang X, Swisher AE, Yeoman PC, Rintoul M, Fleener GB, et al. Ostreid herpesvirus 1 latent infection and reactivation in adult Pacific oysters, Crassostrea gigas. Virus Res. 2024;339:199245. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR218] 218.Eberle R, Maxwell LK, Nicholson S, Black D, Jones-Engel L. Genome sequence variation among isolates of monkey B virus (Macacine alphaherpesvirus 1) from captive macaques. Virology. 2017;508:26–35. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR219] 219.Freifeld AG, Hilliard J, Southers J, Murray M, Savarese B, Schmitt JM, et al. A controlled seroprevalence survey of primate handlers for evidence of asymptomatic herpes B virus infection. J Infect Dis. 1995;171(4):1031–4. [DOI] [PubMed] [Google Scholar]

[CR220] 220.Gaskell R, Dawson S, Radford A, Thiry E. Feline herpesvirus. Vet Res. 2007;38(2):337–54. [DOI] [PubMed] [Google Scholar]

[CR221] 221.Gowthaman V, Kumar S, Koul M, Dave U, Murthy TRGK, Munuswamy P, et al. Infectious laryngotracheitis: Etiology, epidemiology, pathobiology, and advances in diagnosis and control – a comprehensive review. Vet Q. 2020;40(1):140–61. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR222] 222.Holland MS, Mackenzie CD, Bull RW, Silva RF. Latent Turkey herpesvirus infection in lymphoid, nervous, and feather tissues of chickens. Avian Dis. 1998;42(2):292–9. [PubMed] [Google Scholar]

[CR223] 223.Mahalingam R, Clarke P, Wellish M, Dueland AN, Soike KF, Gilden DH, et al. Prevalence and distribution of latent simian varicella virus DNA in monkey ganglia. Virology. 1992;188(1):193–7. [DOI] [PubMed] [Google Scholar]

[CR224] 224.Mekuria ZH, El-Hage C, Ficorilli NP, Washington EA, Gilkerson JR, Hartley CA. Mapping B lymphocytes as major reservoirs of naturally occurring latent equine herpesvirus 5 infection. J Gen Virol. 2017;98(3):461–70. [DOI] [PubMed] [Google Scholar]

[CR225] 225.Mueller NJ, Denner J. Porcine cytomegalovirus/porcine roseolovirus (PCMV/PRV): a threat for xenotransplantation? Xenotransplantation. 2022;29(5):e12775. [DOI] [PubMed] [Google Scholar]

[CR226] 226.Pan L, Li M, Zhang X, Xia Y, Mian AM, Wu H, et al. Establishment of an in vitro model of pseudorabies virus latency and reactivation and identification of key viral latency-associated genes. Viruses. 2023;15(3):808. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR227] 227.Quijano Cardé EM, Soto E. A review of latency in the Alloherpesviridae family. J Fish Dis. 2024;47(12):e14016. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR228] 228.Riaz A, Dry I, Levy CS, Hopkins J, Grey F, Shaw DJ, et al. Ovine herpesvirus-2-encoded microRNAs target virus genes involved in virus latency. J Gen Virol. 2014;95(2):472–80. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR229] 229.Samoilowa S, Giessler KS, Torres CEM, Hussey GS, Allum A, Fux R, et al. Equid herpesvirus-1 distribution in equine lymphoid and neural tissues 70 days post infection. Pathogens. 2021;10(6):707. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR230] 230.Shawky S, Schat KA. Latency sites and reactivation of duck enteritis virus. Avian Dis. 2002;46(2):308–13. [DOI] [PubMed] [Google Scholar]

[CR231] 231.Silvestro C, Bratanich A. The latency related gene of bovine herpesvirus types 1 and 5 and its modulation of cellular processes. Arch Virol. 2016;161(12):3299–308. [DOI] [PubMed] [Google Scholar]

[CR232] 232.Thorsteinsdóttir L, Guðmundsson GÖ, Jensson H, Torsteinsdóttir S, Svansson V. Isolation of equid alphaherpesvirus 3 from a horse in Iceland with equine coital exanthema. Acta Vet Scand. 2021;63(1):6. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR233] 233.van Maanen C. Equine herpesvirus 1 and 4 infections: an update. Veterinary Quarterly. 2002;24(2):57–78. [PubMed] [Google Scholar]

[CR234] 234.Zamana RR, Gattamorta MA, Cruz Ochoa PF, Navas-Suárez PE, Sacristán C, Rossi S, et al. High Occurrence of Chelonid Alphaherpesvirus 5 (ChHV5) in Green Sea Turtles Chelonia mydas with and without Fibropapillomatosis in Feeding Areas of the São Paulo Coast, Brazil. J Aquat Anim Health. 2021;33(4):252–63. [DOI] [PubMed] [Google Scholar]

[CR235] 235.Jin L, Löhr CV, Vanarsdall AL, Baker RJ, Moerdyk-Schauwecker M, Levine C, et al. Characterization of a novel alphaherpesvirus associated with fatal infections of domestic rabbits. Virology. 2008;378(1):13–20. [DOI] [PubMed] [Google Scholar]

[CR236] 236.Slater JD, Borchers K, Thackray AM, Field HJ. The trigeminal ganglion is a location for equine herpesvirus 1 latency and reactivation in the horse. J Gen Virol. 1994;75(8):2007–16. [DOI] [PubMed] [Google Scholar]

[CR237] 237.Stiles J. Feline herpesvirus. Clin Tech Small Anim Pract. 2003;18(3):178–85. [DOI] [PubMed] [Google Scholar]

[CR238] 238.Maidana SS, Delgado F, Vagnoni L, Mauroy A, Thiry E, Romera S. Cattle are a potential reservoir of bubaline herpesvirus 1 (BuHV1). Veterinary Record Open. 2016;3(1):e000162. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR239] 239.Meyer G, Lemaire M, Ros C, Belak K, Gabriel A, Cassart D, et al. Comparative pathogenesis of acute and latent infections of calves with bovine herpesvirus types 1 and 5. Arch Virol. 2001;146(4):633–52. [DOI] [PubMed] [Google Scholar]

[CR240] 240.Ostler JB, Jones C. The bovine herpesvirus 1 latency-reactivation cycle, a chronic problem in the cattle industry. Viruses. 2023;15(2):552. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR241] 241.Mahalingam R, Wellish M, Wolf W, Dueland AN, Cohrs R, Vafai A, et al. Latent varicella-zoster viral DNA in human trigeminal and thoracic ganglia. N Engl J Med. 1990;323(10):627–31. [DOI] [PubMed] [Google Scholar]

[CR242] 242.Thilakarathne DS, Hartley CA, Diaz-Méndez A, Quinteros JA, Fakhri O, Coppo MJC, et al. Latency characteristics in specific pathogen-free chickens 21 and 35 days after intra-tracheal inoculation with vaccine or field strains of infectious laryngotracheitis virus. Avian Pathol. 2020;49(4):369–79. [DOI] [PubMed] [Google Scholar]

[CR243] 243.Borchers K, Wolfinger U, Lawrenz B, Schellenbach A, Ludwig H. Equine herpesvirus 4 DNA in trigeminal ganglia of naturally infected horses detected by direct in situ PCR. J Gen Virol. 1997;78(5):1109–14. [DOI] [PubMed] [Google Scholar]

[CR244] 244.Six A, Banks M, Engels M, Bascuñana CR, Ackermann M. Latency and reactivation of bovine herpesvirus 1 (BHV-1) in goats and of caprine herpesvirus 1 (CapHV-1) in calves. Arch Virol. 2001;146(7):1325–35. [DOI] [PubMed] [Google Scholar]

[CR245] 245.Borchers K, Lieckfeldt D, Ludwig A, Fukushi H, Allen G, Fyumagwa R, et al. Detection of equid herpesvirus 9 DNA in the trigeminal ganglia of a burchell’s zebra from the serengeti ecosystem. J Vet Med Sci. 2008;70(12):1377–81. [DOI] [PubMed] [Google Scholar]

[CR246] 246.Pantry SN, Medveczky PG. Latency, integration, and reactivation of human herpesvirus-6. Viruses. 2017;9(7):194. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR247] 247.Leskowitz RM, Zhou XY, Villinger F, Fogg MH, Kaur A, Lieberman PM, et al. CD4+ and CD8+ T-cell responses to latent antigen EBNA-1 and lytic antigen BZLF-1 during persistent lymphocryptovirus infection of rhesus macaques. J Virol. 2013;87(15):8351–62. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR248] 248.Wei C, Kakazu T, Chuah QY, Tanaka M, Kato G, Sano M. Reactivation of cyprinid herpesvirus 2 (CyHV-2) in asymptomatic surviving goldfish Carassius auratus (L.) under immunosuppression. Fish Shellfish Immunol. 2020;103:302–9. [DOI] [PubMed] [Google Scholar]

[CR249] 249.Thonur L, Russell GC, Stewart JP, Haig DM. Differential transcription of ovine herpesvirus 2 genes in lymphocytes from reservoir and susceptible species. Virus Genes. 2006;32(1):27–35. [DOI] [PubMed] [Google Scholar]

[CR250] 250.Perdrizet UG, Hill JE, Sobchishin L, Singh B, Fernando C, Bollinger TK, et al. Tissue and cellular tropism of Eptesicus fuscus gammaherpesvirus in big brown bats, potential role of pulmonary intravascular macrophages. Vet Pathol. 2024;61(4):550–61. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR251] 251.Mallet C, Cochard J, Leclercq S, Trapp-Fragnet L, Chouteau P, Denesvre C. Hypoxia and HIF-1 trigger Marek’s disease virus reactivation in lymphoma-derived latently infected T Lymphocytes. J Virol. 2022;96(5):e0142721. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR252] 252.Juillard F, Tan M, Li S, Kaye KM. Kaposi’s Sarcoma Herpesvirus Genome Persistence. Front Microbiol. 2016;7. Disponible sur: https://www.frontiersin.org/journals/microbiology/articles/10.3389/fmicb.2016.01149/full. Cité 24 févr 2025. [DOI] [PMC free article] [PubMed]

[CR253] 253.Kennedy PGE, Rovnak J, Badani H, Cohrs RJ. A comparison of herpes simplex virus type 1 and varicella-zoster virus latency and reactivation. J Gen Virol. 2015;96(Pt 7):1581–602. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR254] 254.Speck SH, Ganem D. Viral latency and its regulation: lessons from the γ-Herpesviruses. Cell Host Microbe. 2010;8(1):100–15. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR255] 255.Tempera I, Lieberman PM. Chromatin organization of gammaherpesvirus latent genomes. Biochim Biophys Acta. 2010;1799(3):236–45. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR256] 256.Tarrant-Elorza M, Rossetto CC, Pari GS. Maintenance and replication of the human cytomegalovirus genome during latency. Cell Host Microbe. 2014;16(1):43–54. [DOI] [PubMed] [Google Scholar]

[CR257] 257.Nair V. Latency and tumorigenesis in Marek’s disease. Avian Dis. 2013;57(2s1):360–5. [DOI] [PubMed] [Google Scholar]

[CR258] 258.Morissette G, Flamand L. Herpesviruses and chromosomal Integration. J Virol. 2010;84(23):12100–9. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR259] 259.Arbuckle JH, Medveczky MM, Luka J, Hadley SH, Luegmayr A, Ablashi D, et al. The latent human herpesvirus-6A genome specifically integrates in telomeres of human chromosomes in vivo and in vitro. Proc Natl Acad Sci. 2010;107(12):5563–8. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR260] 260.Kheimar A, Previdelli RL, Wight DJ, Kaufer BB. Telomeres and telomerase: role in marek’s disease virus pathogenesis, integration and tumorigenesis. Viruses. 2017;9(7):173. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR261] 261.Lestou VS, De Braekeleer M, Strehl S, Ott G, Gadner H, Ambros PF. Non-random integration of epstein-barr virus in lymphoblastoid cell lines. Genes Chromosom Cancer. 1993;8(1):38–48. [DOI] [PubMed] [Google Scholar]

[CR262] 262.Ahmed W, Liu ZF. Long Non-Coding RNAs: Novel Players in Regulation of Immune Response Upon Herpesvirus Infection. Frontiers in Immunology. 2018;9. Disponible sur: https://www.frontiersin.org/articles/10.3389/fimmu.2018.00761. Cité 19juill 2023. [DOI] [PMC free article] [PubMed]

[CR263] 263.Cliffe AR, Garber DA, Knipe DM. Transcription of the herpes simplex virus latency-associated transcript promotes the formation of facultative heterochromatin on lytic promoters. J Virol. 2009;83(16):8182–90. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR264] 264.Cullen BR. Viruses and microRNAs: RISCy interactions with serious consequences. Genes Dev. 2011;25(18):1881–94. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR265] 265.Pan D, Flores O, Umbach JL, Pesola JM, Bentley P, Rosato PC, et al. A neuron-specific host MicroRNA targets herpes simplex virus-1 ICP0 expression and promotes latency. Cell Host Microbe. 2014;15(4):446–56. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR266] 266.Grey F. Role of microRNAs in herpesvirus latency and persistence. J Gen Virol. 2015;96(4):739–51. [DOI] [PubMed] [Google Scholar]

[CR267] 267.Morgan RW, Burnside J. Roles of avian herpesvirus microRNAs in infection, latency, and oncogenesis. Biochim Biophys Acta. 2011;1809(11):654–9. [DOI] [PubMed] [Google Scholar]

[CR268] 268.Münz C. Latency and lytic replication in Epstein-Barr virus-associated oncogenesis. Nat Rev Microbiol. 2019;17(11):691–700. [DOI] [PubMed] [Google Scholar]

[CR269] 269.Chida Y, Mao X. Does psychosocial stress predict symptomatic herpes simplex virus recurrence? A meta-analytic investigation on prospective studies. Brain Behav Immun. 2009;23(7):917–25. [DOI] [PubMed] [Google Scholar]

[CR270] 270.El Hayderi L, Delvenne P, Rompen E, Senterre JM, Nikkels AF. Herpes simplex virus reactivation and dental procedures. Clin Oral Investig. 2013;17(8):1961–4. [DOI] [PubMed] [Google Scholar]

[CR271] 271.Padgett DA, Sheridan JF, Dorne J, Berntson GG, Candelora J, Glaser R. Social stress and the reactivation of latent herpes simplex virus type 1. Proc Natl Acad Sci. 1998;95(12):7231–5. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR272] 272.Roizman B, Whitley RJ. An inquiry into the molecular basis of HSV latency and reactivation. Annu Rev Microbiol. 2013;67:355–74. [DOI] [PubMed] [Google Scholar]

[CR273] 273.Suzich JB, Cliffe AR. Strength in diversity: Understanding the pathways to herpes simplex virus reactivation. Virology. 2018;522:81–91. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR274] 274.Tien YT, Akbar H, Jarosinski KW. Temperature-induced reactivation of Marek’s disease virus-transformed T cells ex vivo. Front Vet Sci. 2023;10:1145757. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR275] 275.Du T, Zhou G, Roizman B. HSV-1 gene expression from reactivated ganglia is disordered and concurrent with suppression of latency-associated transcript and miRNAs. Proc Natl Acad Sci. 2011;108(46):18820–4. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR276] 276.Kim JY, Mandarino A, Chao MV, Mohr I, Wilson AC. Transient reversal of episome silencing precedes VP16-dependent transcription during reactivation of latent HSV-1 in Neurons. PLoS Pathog. 2012;8(2):e1002540. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR277] 277.Hale AE, Moorman NJ. The ends dictate the means: promoter switching in herpesvirus gene expression. Annu Rev Virol. 2021;8:201–18. [DOI] [PubMed] [Google Scholar]

[CR278] 278.Jia Z, Zhang D, Zhu L, Xue J. Animal models of human herpesvirus infection. Anim Models Exp Med. 2025;8(4):615–28. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR279] 279.Petton B, Pernet F, Robert R, Boudry P. Temperature influence on pathogen transmission and subsequent mortalities in juvenile Pacific oysters Crassostrea gigas. Aquacult Environ Interact. 2013;3(3):257–73. [Google Scholar]

[CR280] 280.Rybak-Wolf A, Wyler E, Pentimalli TM, Legnini I, Oliveras Martinez A, Glažar P, et al. Modelling viral encephalitis caused by herpes simplex virus 1 infection in cerebral organoids. Nat Microbiol. 2023;8(7):1252–66. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR281] 281.Pourmaleki M, Jones CJ, Mellinghoff SD, Greenstein BD, Kumar P, Foronda M, et al. Multiplexed spatial profiling of hodgkin reed-sternberg cell neighborhoods in classic hodgkin lymphoma. Clin Cancer Res. 2024;30(17):3881–93. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR282] 282.Tombácz D, Balázs Z, Csabai Z, Moldován N, Szűcs A, Sharon D, et al. Characterization of the dynamic transcriptome of a herpesvirus with long-read single molecule real-time sequencing. Sci Rep. 2017;7(1):43751. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR283] 283.Rosani U, Bortoletto E, Zhang X, Huang BW, Xin LS, Krupovic M, et al. Long-read transcriptomics of Ostreid herpesvirus 1 uncovers a conserved expression strategy for the capsid maturation module and pinpoints a mechanism for evasion of the ADAR-based antiviral defence. Virus Evol. 2024;10(1):088. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR284] 284.Singh RK, Vangala R, Torne AS, Bose D, Robertson ES. Epigenetic and epitranscriptomic regulation during oncogenic γ-herpesvirus infection. Front Microbiol. 2024;15:1484455. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR285] 285.Sun A, Zhu X, Liu Y, Wang R, Yang S, Teng M, et al. Transcriptome-wide N6-methyladenosine modification profiling of long non-coding RNAs during replication of Marek’s disease virus in vitro. BMC Genomics. 2021;22(1):296. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR286] 286.Feng Z, Zhou F, Tan M, Wang T, Chen Y, Xu W, et al. Targeting m6A modification inhibits herpes virus 1 infection. Genes Dis. 2022;9(4):1114–28. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR287] 287.Gao C, Chen L, Tang SB, Long QY, He JL, Zhang NA, et al. The epigenetic landscapes of histone modifications on HSV-1 genome in human THP-1 cells. Antiviral Res. 2020;176:104730. [DOI] [PubMed] [Google Scholar]

[CR288] 288.Volkening JD, Spatz SJ, Ponnuraj N, Akbar H, Arrington JV, Vega-Rodriguez W, et al. Viral proteogenomic and expression profiling during productive replication of a skin-tropic herpesvirus in the natural host. PLoS Pathog. 2023;19(6):e1011204. [DOI] [PMC free article] [PubMed] [Google Scholar]

[CR289] 289.Jumper J, Evans R, Pritzel A, Green T, Figurnov M, Ronneberger O, et al. Highly accurate protein structure prediction with AlphaFold. Nature. 2021;596(7873):583–9. [DOI] [PMC free article] [PubMed] [Google Scholar]

PERMALINK

Herpesviruses: overview of systematics, genomic complexity and life cycle

Aurélie Dotto-Maurel

Isabelle Arzul

Benjamin Morga

Germain Chevignon

Abstract

Background

Fig. 1.

Herpesviruses virion structure

Fig. 2.

Envelope

Tegument

Capsid

Core

Towards a better characterization of herpesvirus particle structure

Herpesvirales clade

Fig. 3.

Orthoherpesviridae

Alphaherpesvirinae

Table 1.

Betaherpesvirinae

Table 2.

Gammaherpesvirinae

Table 3.

Malacoherpesviridae

Table 4.

Alloherpesviridae

Table 5.

Perspective on Herpesvirus classification

Genomic architecture of Herpesvirales

Table 6.

Orthoherpesviridae

Alphaherpesvirinae

Betaherpesvirinae

Gammaherpesvirinae

Malacoherpesviridae

Alloherpesviridae

Insights into the role of herpesvirus genomic architecture

Genomic isomerization of Herpesvirales and its phenotypic implications

Herpesviruses life cycle

Lytic phase

Fig. 4.

Persistent phase

Maintenance of the viral genome during latency

Fig. 5.

Regulation of viral genes expression during latency

Fig. 6.

Reactivation

Perspectives and insights into herpesvirus life cycle comprehension

Conclusion

Acknowledgements

Abbreviations

Authors’ contributions

Funding

Data availability

Declarations

Ethics approval and consent to participate

Consent for publication

Competing interests

Footnotes

Contributor Information

References

Associated Data

Data Availability Statement

ACTIONS

PERMALINK

RESOURCES

Similar articles

Cited by other articles

Links to NCBI Databases