Baculoviridae - v201911


Baculoviridae - v201911

Baculoviridae

Robert L. Harrison, Elisabeth A. Herniou, Johannes A. Jehle​, David A. Theilmann4, John P. Burand, James J. Becnel, Peter J. Krell, Monique M. van Oers​, Joseph D. Mowery​ and Gary R. Bauchan

Chapter contents

Posted June 2018

Baculoviridae: The family

Member taxa

Supporting information

Citation

A summary of this ICTV Report chapter has been published as an ICTV Virus Taxonomy Profile article in the Journal of General Virology, and should be cited when referencing this online chapter as follows:

Harrison, R.L., Herniou, E.A., Jehle, J.A., Theilmann, D.A., Burand, J.P., Becnel, J.J., Krell, P.J., M. van Oers, M., Mowery, J.D., Bauchan, G.R., and ICTV Report Consortium. 2019, ICTV Virus Taxonomy Profile: BaculoviridaeJournal of General Virology, 99: 11851186.

Summary

The Baculoviridae is a family of large, insect-specific viruses with circular dsDNA genomes ranging from 80 to 180 kbp.  Virions consist of enveloped, rod-shaped nucleocapsids and are embedded in distinctive occlusion bodies measuring 0.15–5 µm.  The occlusion bodies consist of a matrix composed of a single viral protein expressed at high levels during infection.  Members of this family infect exclusively larvae of insect orders Lepidoptera, Hymenoptera and Diptera.  Some members have been developed as biopesticides for controlling insect pests and as vectors for recombinant protein expression. 

Table 1.Baculoviridae. Characteristics of the family Baculoviridae.

Characteristic

Description

Typical member

Autographa californica multiple nucleopolyhedrovirus C6 (L22858), species Autographa californica multiple nucleopolyhedrovirus, genus Alphabaculovirus

Virion

One or two distinct types of virions consisting of enveloped, rod-shaped nucleocapsids, 30–60 nm × 250–300 nm, containing >20 proteins

Genome

A single covalently-closed circular dsDNA molecule of 80–180 kbp encoding 100–200 proteins

Replication

Nuclear, with nucleocapsids assembled in the nucleus and enveloped either (a) in the nucleus or mixed nucleoplasm and cytoplasm, or (b) upon budding through the plasma membrane

Translation

From mRNAs transcribed from viral DNA

Host range

Larval-stage insects of orders Diptera, Hymenoptera, and Lepidoptera

Taxonomy

Four genera with 76 species

Virion

Morphology

One or two virion phenotypes are involved in baculovirus infections. Infection is initiated in the gut epithelium by virions contained within a crystalline protein occlusion body (OB) which may be polyhedral in shape and containing many virions (members of the genera Alphabaculovirus, Gammabaculovirus and Deltabaculovirus); or ovocylindrical and containing only one, or rarely two or more, virions (members of the genus Betabaculovirus). Virions within occlusions, referred to as occlusion-derived virus (ODV), consist of one or more rod-shaped nucleocapsids that have a distinct structural polarity and are enclosed within an envelope (Figure 1.Baculoviridae). For ODV, nucleocapsid envelopment occurs within the nucleus (members of the genus Alphabaculovirus) or in the nuclear-cytoplasmic milieu after loss of the nuclear membrane (members of the genus Betabaculovirus). Nucleocapsids average 30–60 nm in diameter and 250–300 nm in length. Spike-like structures (peplomers) have not been reported on envelopes of ODV. Virions of the second phenotype (termed budded virus or BV; Figure 1.Baculoviridae) are generated when nucleocapsids bud through the plasma membrane at the surface of infected cells. BVs typically contain a single nucleocapsid. Their envelopes are derived from the cellular plasma membrane and characteristically appear as a loose-fitting membrane that contains an envelope fusion glycoprotein (EFP), which forms peplomers usually observed at one end of the virion (see “Proteins”, below).

Figure 1.Baculoviridae. Baculovirus virions and nucleocapsids. (Top) The two baculovirus virion phenotypes are illustrated as diagrams with shared and phenotype-specific components.  (Bottom) Transmission electron micrographs of an occlusion-derived virion of Autographa californica multiple nucleopolyhedrovirus (bottom left), a negatively-stained nucleocapsid of Autographa californica multiple nucleopolyhedrovirus (center), and a budded virion of Lymantria dispar multiple nucleopolyhedrovirus (bottom right).  Nucleocapsid and budded virion micrographs courtesy of J. R. Adams (USDA).

Physicochemical and physical properties

ODV buoyant density in CsCl is 1.18–1.25 g cm−3, and that of the nucleocapsid is 1.47 g cm−3. BV buoyant density in sucrose is 1.17–1.18 g cm−3 (Summers and Volkman 1976). Virions of both phenotypes are sensitive to organic solvents and detergents. ODV and BV are marginally sensitive to heat and inactivated at extreme pH values (Gudauskas and Canerday 1968, Knittel and Fairbrother 1987).

Nucleic acid

Nucleocapsids contain one molecule of circular, supercoiled dsDNA of 80–180 kbp (Figure 1.Baculoviridae, Figure 2.Baculoviridae).

Proteins

Genomic analyses suggest that baculoviruses encode approximately 100–200 proteins.  Proteomics analyses to date indicate that virions may contain as few as 23 and as many as 73 different polypeptides (Deng et al., 2007, Zhang et al., 2015). Nucleocapsids from both virion phenotypes (ODV and BV) contain a major capsid protein (VP39), a basic DNA-binding protein (P6.9) complexed with the viral genome, and at least 2–3 additional proteins. Two different EFPs have been identified to date.  The EFP GP64 is present in a group of alphabaculoviruses that include Autographa californica multiple nucleopolyhedrovirus (AcMNPV) and close relatives (Blissard and Wenz 1992). Most of the alphabaculoviruses and betabaculoviruses encode and appear to utilize EFPs known as F proteins, which are homologs of the Ld130 protein from Lymantria dispar multiple nucleopolyhedrovirus (LdMNPV) (Pearson et al., 2001) and the Se8 protein from Spodoptera exigua multiple nucleopolyhedrovirus (SeMNPV) (IJkel et al., 2000). The deltabaculovirus Culex nigripalpus nucleopolyhedrovirus also encodes an EFP with fusogenic activity (Wang et al., 2017). Several ODV envelope proteins have been identified. Eight ODV proteins, including P74 (Faulkner et al., 1997), PIF-1 (Kikhno et al., 2002), PIF-2 (Pijlman et al., 2003), PIF-3 (Ohkawa et al., 2005), AC96 (PIF-4) (Fang et al., 2009), ODV-E56 (PIF-5) (Harrison et al., 2010), AC68 (PIF-6) (Nie et al., 2012), AC110 (PIF-7) (Jiantao et al., 2016), and AC83 (PIF-8) (Javed et al., 2017) are essential for oral infectivity of ODV. The major protein of the OB matrix is a virus-encoded polypeptide of 25–33 kDa. This protein is called polyhedrin for nucleopolyhedroviruses (the common name used for the alpha-, delta- and gammabaculoviruses) and granulin for granuloviruses (betabaculoviruses) (Rohrmann 1986). The OB is often surrounded by an envelope that contains at least one major protein (Whitt and Manning 1988). The polyhedrin protein of deltabaculoviruses is serologically and genetically unrelated to OB proteins of the alpha-, beta- and gammabaculoviruses (Perera et al., 2006).

Lipids

Lipids are present in the envelopes of ODV and BV. Lipid composition differs between the two virion phenotypes (Braunagel and Summers 1994).

Carbohydrates

Carbohydrates are present as glycoproteins and glycolipids.

Genome organization and replication

Circular genomic DNA is infectious, suggesting that after cellular entry and uncoating, no virion-associated proteins are essential for infection (Burand et al., 1980, Carstens et al., 1980). Thirty-eight gene homologs, the so-called baculovirus core genes, are shared among alpha-, beta-, gamma- and deltabaculoviruses (Javed et al., 2017, Garavaglia et al., 2012) (Figure 2.Baculoviridae). These conserved genes are involved in various functions, including DNA replication, late gene transcription and virion structure. In some cases, larger genome sizes may result from the presence of families of repeated genes (Hayakawa et al., 1999). Transcription of baculovirus genes is temporally regulated, and two main classes of genes are recognized: early and late. Late genes may be further subdivided into late and very late genes. Gene classes (early, late and very late) are not clustered on the baculovirus genome, and both strands of the genome are involved in coding functions. Early genes are transcribed by host RNA polymerase II, whereas late and very late genes are transcribed by an alpha-amanitin-resistant viral RNA polymerase (Huh and Weaver 1990). RNA splicing occurs, but appears to be rare (Chisholm and Henner 1988, Pearson and Rohrmann 1997). Transient early and late gene transcription and DNA replication studies suggest that at least three virus-encoded proteins regulate early gene transcription (Guarino and Summers 1986, Kovacs et al., 1991, Lu and Carstens 1993, Yoo and Guarino 1994), whereas approximately 20 viral encoded proteins known as late expression factors (LEFs) are necessary for late gene transcription (Rapp et al., 1998, Huijskens et al., 2004). Of the approximately 20 LEFs, half appear to be involved in DNA replication (Lu and Miller 1995). Late gene transcription initiates at the second adenine of a conserved 5′-TAAG-3′ promoter motif, which is an essential core element of the baculovirus late promoter (Chen et al., 2013). Putative replication origins consist of repeated sequences found at multiple locations within the baculovirus genome (Pearson et al., 1992, Hilton and Winstanley 2007). These sequences, termed homologous repeat (hr) regions, do not appear to be highly conserved among different baculovirus species. Single copy, non-hr putative replication origins have also been identified (Kool et al., 1994). DNA replication is required for late gene transcription. Most virion structural proteins are encoded by late genes. While transcription of late and very late genes appears to begin immediately after DNA replication, some very late genes that encode occlusion body-specific proteins are transcribed at extremely high levels at a later time (Thiem and Miller 1990). BV production occurs primarily during the late phase, and occlusion body production occurs during the very late phase.

Figure 2.Baculoviridae. Genome map of Autographa californica multiple nucleopolyhedrovirus isolate C6, the representative isolate of the genus Alphabaculovirus type species Autographa californica multiple nucleopolyhedrovirus.  The genome is illustrated with locations and orientations of annotated ORFs (arrows). ORFs corresponding to the core genes of members of the family Baculoviridae, and ORFs conserved among alpha-and betabaculoviruses are indicated.  Locations of homologous repeat (hr) sequences are also shown, as well as the exons and intron of the ie-0 transcription unit. 

In infected animals, viral replication begins in the insect midgut. Following ingestion, OBs are solubilized in the gut lumen, releasing the ODVs, which are thought to enter the target epithelial cells via fusion with the plasma membrane at the cell surface (Kawanishi et al., 1972). In lepidopteran insects, viral entry into midgut cells occurs in an alkaline environment, up to pH 12. Infection of the midgut is required for initiation of infection in the animal. Although the virus is believed to undergo one round of replication in the midgut epithelium prior to transmission of infection to secondary tissues within the hemocoel, a mechanism for direct movement from the midgut to the hemocoel has also been proposed (Granados and Lawler 1981, Washburn et al., 1999, Washburn et al., 2003).

DNA replication takes place in the nucleus. In betabaculovirus-infected cells, the integrity of the nuclear membrane is lost during the replication process (Walker et al., 1982, Federici and Stern 1990). With some baculoviruses, replication is restricted to the gut epithelium and progeny virions become enveloped and occluded within these cells, and may be shed into the gut lumen with sloughed epithelium, or released upon death of the host (Federici and Stern 1990). In other baculoviruses, the infection is transmitted to internal organs and tissues (Keddie et al., 1989). The second virion phenotype, BV, which buds from the basolateral membrane of infected gut cells, is required for transmission of the infection into the hemocoel. Infected fat body cells are the primary location of occluded virus production in lepidopteran insects. Occluded virus matures within nuclei of infected cells for alpha-, gamma- and deltabaculoviruses, and within the nuclear-cytoplasmic milieu for betabaculoviruses. OBs containing infectious ODV virions are released upon death, and usually liquefaction, of the host. 

Antigenicity

Antigenic determinants that cross-react between different baculoviruses exist on virion proteins and on the major OB polypeptide: polyhedrin or granulin (Summers et al., 1978, Volkman 1983). Neutralizing antibodies react with the major surface glycoprotein of BVs (Volkman et al., 1984).

Biology

Baculoviruses have been isolated from insects only - primarily from insects of the order Lepidoptera, but also from the orders Hymenoptera and Diptera. Transmission naturally occurs (i) horizontally by contamination with OBs of food, egg surfaces, etc. (Hamm and Young 1974, Young and Yearian 1986); (ii) vertically within the egg either from infected female or male adults (Doane 1969); or experimentally; (iii) by injection of intact hosts with BVs; or (iv) by infection or transfection of cell cultures. Typically, the infection process in a permissive insect host requires approximately one week, and, as an end result, the diseased insect liquefies, releasing infectious occlusion bodies into the environment. OBs represent an environmentally stable form of the virus with increased resistance to chemical and physical decay as well as inactivation by UV light.  Persistent, asymptomatic infections have also been documented (Myers and Cory 2016).

Genus demarcation criteria

Genera of Baculoviridae are distinguished on the basis of phylogeny, genome characteristics (especially gene content), host range and OB morphology (Jehle et al., 2006a).

Derivation of names

Alpha, Beta, Gamma, Delta: Greek letters α, β, γ, and δ, the first four letters of the Greek alphabet.

Baculo-: from baculum, meaning “rod” in latin, referring to the morphology of the nucleocapsid.

Granulo-: from “granule” and “granulosis”, referring to the relatively small size of OBs and their granular appearance in betabaculovirus-infected cells.

Nucleopolyhedro-: from “nuclear polyhedrosis” and “polyhedron”, referring to the multifaceted appearance of OBs in the nuclei of infected cells. 

Phylogenetic relationships

Phylogenetic analysis based on the 38 baculovirus core genes shows that the family comprises four monophyletic groups (Figure 3.Baculoviridae), which can also be discriminated based on the orders of their insect hosts and on their morphology. Baculoviruses are classified into the four genera Alphabaculovirus, Betabaculovirus, Gammabaculovirus and Deltabaculovirus.

Figure 3.Baculoviridae. Phylogeny of family Baculoviridae. The maximum likelihood tree, based on the alignment of 38 core gene amino acid sequences, shows the relationships of the species for which a completely annotated genome of a representative isolate was available at time of analysis. Amino acid sequences of the 38 core genes were aligned individually by MUSCLE and concatenated, and phylogeny was inferred from the concatenated alignments with the Le and Gascuel (LG) substitution model using RAxML with 100 bootstrap replicates. Colored circles indicate genus assignments for each virus; unclassified viruses are indicated by open circles. This phylogenetic tree and corresponding sequence alignment are available to download from the Resources page.

Similarity with other taxa

Members of the family Baculoviridae share structural, genetic and biological characters with viruses of family Nudiviridae, which had been formerly classified as “non-occluded” baculoviruses. The nudiviruses share at least 20 core genes with baculoviruses (Wang et al., 2011).  Baculoviruses are also similar to the salivary gland hypertrophy viruses of Hytrosaviridae, and share at least 12 core genes with members of this family (Jehle et al., 2013). 


Genus: Alphabaculovirus

Distinguishing features 

OBs measure 0.15 to 5 µm in size and are generally polyhedral, although morphology may vary (Adams and McClintock 1991).  They mature within the nuclei of infected cells and characteristically contain many enveloped virions (Figure 1.Alphabaculovirus). The occluded virions are packaged with either single or multiple nucleocapsids within a single viral envelope. Members of some virus species manifest both phenotypes. A number of genes that influence nucleocapsid packaging are known, but in members of some species, packaging may be variable (Rohrmann 2014, Yang et al., 2014). Single (S) and multiple (M) designations in common names have been retained for species where variability has not been reported and for distinct viruses that would otherwise have identical designations under the current nomenclature. During viral entry, nucleocapsids are transported through the nuclear membrane and into the nucleus (Au et al., 2016), where uncoating and viral replication occur.

Figure 1.Alphabaculovirus. Occlusion bodies of baculoviruses in the genus Alphabaculovirus. Scanning electron micrographs of Autographa californica multiple nucleopolyhedrovirus (A) and Operophtera brumata nucleopolyhedrovirus (B) occlusion bodies are shown, along with transmission electron micrographs of Autographa californica multiple nucleopolyhedrovirus (C) and Trichoplusia ni single nucleopolyhedrovirus (D) occlusion bodies containing occlusion-derived virus consisting of multiple and single nucleocapsids per envelope, respectively.  Scale bars: (A, B) 2 µm; (C, D) 0.5 µm.  Micrographs in panels (A) and (D) courtesy of J. R. Adams (USDA).

Virion

Morphology

Nucleocapsid length appears to be proportional to genome size (Fraser 1986).

Genome organization and replication

In addition to the family Baculoviridae core genes, the alphabaculoviruses and betabaculoviruses appear to share an additional 23 homologs (Garavaglia et al., 2012).

Biology

Species of this genus have been isolated only from the insect order Lepidoptera. 

Species demarcation criteria

Traditionally, species distinctions have been broadly based on host range and specificity, DNA restriction profiles, DNA sequences from various regions of the genome, and predicted protein sequence similarities.  More recently, species demarcation criteria for alpha- and betabaculoviruses have been proposed that rely upon pairwise nucleotide distances estimated with the Kimura-2-parameter substitution model from partial sequences of three conserved baculovirus genes: late expression factor-8 (lef-8) and late expression factor-9 (lef-9), which encode subunits of the baculovirus RNA polymerase, and polyhedrin/granulin (polh/gran), which encodes the viral occlusion body matrix protein (Jehle et al., 2006b). If nucleotide distances between two viruses at these loci are less than 0.015 substitutions/site, the two baculoviruses are considered to belong to the same species. If nucleotide distances between two viruses are greater than 0.05 substitutions/site, the viruses are considered to belong to different species. If the nucleotide distances lie between 0.015 and 0.050 substitutions/site, additional characteristics of the two viruses (i.e. host range) must be considered to make a decision about their taxonomic status. The proposed criteria were originally based on an alignment of sequences from 117 separate baculovirus isolates and the phylogeny inferred from this alignment. Researchers have applied this criterion to other isolates to identify many new baculovirus species and variants of currently recognized species. A more recent examination of pairwise nucleotide distances for all 38 baculovirus core genes among 172 completely sequenced baculovirus genomes has confirmed the current species classification based on pairwise distances for lef-8lef-9, and polh loci (Wennmann et al., 2018).

Member species

Species Virus name(s) Exemplar isolate Exemplar accession number Exemplar RefSeq number Available sequence Other isolates Other isolate accession numbers Virus abbreviation Isolate abbreviation
Adoxophyes honmai nucleopolyhedrovirus Adoxophyes honmai nucleopolyhedrovirus ADN001 AP006270 NC_004690 Complete genome AdhoNPV
Agrotis ipsilon multiple nucleopolyhedrovirus Agrotis ipsilon multiple nucleopolyhedrovirus Illinois EU839994 NC_011345 Complete genome AgipMNPV
Agrotis segetum nucleopolyhedrovirus A Agrotis segetum nucleopolyhedrovirus A Polish DQ123841 NC_007921 Complete genome AgseNPV-A
Agrotis segetum nucleopolyhedrovirus B Agrotis segetum nucleopolyhedrovirus B English KM102981 NC_025960 Complete genome AgseNPV-B
Antheraea pernyi nucleopolyhedrovirus Antheraea pernyi nucleopolyhedrovirus Liaoning DQ486030 NC_008035 Complete genome AnpeNPV
Anticarsia gemmatalis multiple nucleopolyhedrovirus Anticarsia gemmatalis multiple nucleopolyhedrovirus 2D DQ813662 NC_008520 Complete genome AgMNPV
Anticarsia gemmatalis multiple nucleopolyhedrovirus Anticarsia gemmatalis multiple nucleopolyhedrovirus 37 KR815466 AgMNPV
Autographa californica multiple nucleopolyhedrovirus Autographa californica multiple nucleopolyhedrovirus C6 L22858 NC_001623 Complete genome AcMNPV
Autographa californica multiple nucleopolyhedrovirus Plutella xylostella multiple nucleopolyhedrovirus CL3 DQ457003 PlxyMNPV
Bombyx mori nucleopolyhedrovirus Bombyx mori nucleopolyhedrovirus T3 L33180 NC_001962 Complete genome BmNPV
Bombyx mori nucleopolyhedrovirus Bombyx mandarina nucleopolyhedrovirus S1 FJ882854 BomaNPV
Buzura suppressaria nucleopolyhedrovirus Buzura suppressaria nucleopolyhedrovirus Hubei KF611977 NC_023442 Complete genome BuzuNPV
Catopsilia pomona nucleopolyhedrovirus Catopsilia pomona nucleopolyhedrovirus 416 KU565883 NC_030240 Complete genome CapoNPV
Choristoneura fumiferana DEF multiple nucleopolyhedrovirus Choristoneura fumiferana DEF multiple nucleopolyhedrovirus AY327402 NC_005137 Complete genome CfDEFMNPV
Choristoneura fumiferana multiple nucleopolyhedrovirus Choristoneura fumiferana multiple nucleopolyhedrovirus Ireland AF512031 NC_004778 Complete genome CfMNPV
Choristoneura murinana nucleopolyhedrovirus Choristoneura murinana nucleopolyhedrovirus Darmstadt KF894742 NC_023177 Complete genome ChmuNPV
Choristoneura rosaceana nucleopolyhedrovirus Choristoneura rosaceana nucleopolyhedrovirus NB_1 KC961304 NC_021924 Complete genome ChroNPV
Chrysodeixis chalcites nucleopolyhedrovirus Chrysodeixis chalcites nucleopolyhedrovirus AY864330 NC_007151 Complete genome ChchNPV
Chrysodeixis includens nucleopolyhedrovirus Pseudoplusia includens single nucleopolyhedrovirus IE KJ631622 NC_026268 Complete genome PsinSNPV
Clanis bilineata nucleopolyhedrovirus Clanis bilineata nucleopolyhedrovirus DZ1 DQ504428 NC_008293 Complete genome ClbiNPV
Ectropis obliqua nucleopolyhedrovirus Ectropis obliqua nucleopolyhedrovirus A1 DQ837165 NC_008586 Complete genome EcobNPV
Epiphyas postvittana nucleopolyhedrovirus Epiphyas postvittana nucleopolyhedrovirus AY043265 NC_003083 Complete genome EppoNPV
Euproctis pseudoconspersa nucleopolyhedrovirus Euproctis pseudoconspersa nucleopolyhedrovirus Hangzhou FJ227128 NC_012639 Complete genome EupsNPV
Helicoverpa armigera nucleopolyhedrovirus Helicoverpa armigera nucleopolyhedrovirus G4 AF271059 NC_002654 Complete genome HearNPV
Helicoverpa armigera nucleopolyhedrovirus Helicoverpa armigera nucleopolyhedrovirus C1 AF303045 HearNPV
Helicoverpa armigera nucleopolyhedrovirus Helicoverpa armigera nucleopolyhedrovirus NNg1 AP010907 HearNPV
Hemileuca species nucleopolyhedrovirus Hemileuca sp. nucleopolyhedrovirus MEM KF158713 NC_021923 Complete genome HespNPV-MEM
Hyphantria cunea nucleopolyhedrovirus Hyphantria cunea nucleopolyhedrovirus N9 AP009046 NC_007767 Complete genome HycuPV
Lambdina fiscellaria nucleopolyhedrovirus Lambdina fiscellaria nucleopolyhedrovirus GR15 KP752043 NC_026922 Complete genome LafiNPV
Leucania separata nucleopolyhedrovirus Leucania separata nucleopolyhedrovirus AH1 AY394490 NC_008348 Complete genome LeseNPV
Lonomia obliqua nucleopolyhedrovirus Lonomia obliqua multiple nucleopolyhedrovirus SP/2000 KP763670 NC_043520 Complete genome LoobNPV-SP/2000
Lymantria dispar multiple nucleopolyhedrovirus Lymantria dispar multiple nucleopolyhedrovirus 5/6 AF081810 NC_001973 Complete genome LdMNPV
Lymantria xylina nucleopolyhedrovirus Lymantria xylina multiple nucleopolyhedrovirus 5 GQ202541 NC_013953 Complete genome LyxyMNPV
Mamestra brassicae multiple nucleopolyhedrovirus Mamestra brassicae multiple nucleopolyhedrovirus K1 JQ798165 NC_023681 Complete genome MbMNPV
Mamestra configurata nucleopolyhedrovirus A Mamestra configurata nucleopolyhedrovirus A 90/2 U59461 NC_003529 Complete genome MacoNPV-A
Mamestra configurata nucleopolyhedrovirus A Mamestra configurata nucleopolyhedrovirus A 90/4 AF539999 MacoNPV-A
Mamestra configurata nucleopolyhedrovirus B Mamestra configurata nucleopolyhedrovirus B 96B AY126275 NC_004117 Complete genome MacoNPV-B
Maruca vitrata nucleopolyhedrovirus Maruca vitrata nucleopolyhedrovirus MV-8 EF125867 NC_008725 Complete genome MaviNPV
Mythimna unipuncta nucleopolyhedrovirus Mythimna unipuncta nucleopolyhedrovirus #7 MF375894 NC_043530 Complete genome MyunNPV#7
Operophtera brumata nucleopolyhedrovirus Operophtera brumata nucleopolyhedrovirus MA MF614691 NC_040621 Complete genome OpbuNPV-MA
Orgyia leucostigma nucleopolyhedrovirus Orgyia leucostigma nucleopolyhedrovirus CFS-77 EU309041 NC_010276 Complete genome OrleNPV
Orgyia pseudotsugata multiple nucleopolyhedrovirus Orgyia pseudotsugata multiple nucleopolyhedrovirus U75930 NC_001875 Complete genome OpMNPV
Oxyplax ochracea nucleopolyhedrovirus Oxyplax ochracea nucleopolyhedrovirus 435 MF143631 NC_043529 Complete genome OxocNPV-435
Peridroma saucia nucleopolyhedrovirus Peridroma species nucleopolyhedrovirus GR167 KM009991 NC_024625 Complete genome PespNPV-GR167
Perigonia lusca nucleopolyhedrovirus Perigonia lusca single nucleopolyhedrovirus KM596836 NC_027923 Complete genome PeluSNPV
Spodoptera exigua multiple nucleopolyhedrovirus Spodoptera exigua multiple nucleopolyhedrovirus US1 AF169823 NC_002169 Complete genome SeMNPV
Spodoptera frugiperda multiple nucleopolyhedrovirus Spodoptera frugiperda multiple nucleopolyhedrovirus 3AP2 EF035042 NC_009011 Complete genome SfMNPV
Spodoptera littoralis nucleopolyhedrovirus Spodoptera littoralis nucleopolyhedrovirus AN1956 JX454574 NC_038369 Complete genome SpliNPV
Spodoptera litura nucleopolyhedrovirus Spodoptera litura nucleopolyhedrovirus G2 AF325155 NC_003102 Complete genome SpltNPV
Sucra jujuba nucleopolyhedrovirus Sucra jujuba nucleopolyhedrovirus 473 KJ676450 NC_028636 Complete genome SujuNPV
Thysanoplusia orichalcea nucleopolyhedrovirus Thysanoplusia orichalcea nucleopolyhedrovirus p2 JX467702 NC_019945 Complete genome ThorNPV
Trichoplusia ni single nucleopolyhedrovirus Trichoplusia ni single nucleopolyhedrovirus DQ017380 NC_007383 Complete genome TnSNPV
Wiseana signata nucleopolyhedrovirus Wiseana signata nucleopolyhedrovirus AF016916 NC_038370 Partial genome WisiNPV
Virus names, the choice of exemplar isolates, and virus abbreviations, are not official ICTV designations.

Related, unclassified viruses

Virus name

Accession number

Virus abbreviation

Apocheima cinerarium nucleopolyhedrovirus

FJ914221

ApciNPV

Cerapteryx graminis nucleopolyhedrovirus

HQ603182, HQ603183, HQ603184

CegrNPV

Condylorrhiza vestigialis multiple nucleopolyhedrovirus

KJ631623

CoveMNPV

Dasychira pudibunda nucleopolyhedrovirus

KP747440

DapuNPV-ML1

Helicoverpa armigera multiple nucleopolyhedrovirus

EU730893

HearMNPV

Lacanobia oleracea nucleopolyhedrovirus

LaolNPV

Leucoma salicis multiple nucleopolyhedrovirus

AY729808, AY729809, AY729810

LesaMNPV

Malacosoma californicum pluviale nucleopolyhedrovirus

AF535137

McplNPV

Malacosoma neustria nucleopolyhedrovirus

AY519243, AY519244, AY519245

ManeNPV

Orgyia pseudotsugata single nucleopolyhedrovirus

AY895150, AY895151, AY895152, AY895153

OpSNPV

Panolis flammea nucleopolyhedrovirus

D00437

PaflNPV

Philosamia cynthia nucleopolyhedrovirus

JX404026

PhcyNPV

Rachiplusia ou multiple nucleopolyhedrovirus

AY145471

RoMNPV-R1

Spodoptera litura nucleopolyhedrovirus II

EU780426

SpltNPV-II

Urbanus proteus nucleopolyhedrovirus

KR011717

UrprNPV-Southern Brazil

Virus names and virus abbreviations, are not official ICTV designations.

Genus: Betabaculovirus

Distinguishing features 

OBs are generally ovocylindrical in shape, measure approximately 0.12 × 0.50 µm, and characteristically contain one virion (Tanada and Hess 1991) (Figure 1.Betabaculovirus). Each ODV virion typically contains a single nucleocapsid within a single envelope. Occluded virions may mature among nuclear-cytoplasmic cellular contents after loss of the nuclear membrane of infected cells. Uncoating is thought to occur by a mechanism in which viral DNA is extruded into the nucleus through the nuclear pore while the capsid remains in the cytoplasm (Summers 1971, Au et al., 2013).

Figure 1.Betabaculovirus. Occlusion bodies of baculoviruses in the genus Betabaculovirus.  Scanning electron micrograph of Spodoptera frugiperda granulovirus occlusion bodies (A, courtesy of J. R. Adams), and transmission electron micrographs of Mythimna unipuncta granulovirus B occlusion bodies (B and C), showing their unique ovocylindrical shape. Scale bars: (A) 2 µm; (B, C) 100 nm. 

Virion

See discussion under family description.

Genome organization and replication 

See discussion under family description.

Biology

Members of this genus have been isolated only from the insect order Lepidoptera.

Species demarcation criteria

Traditionally, species distinctions have been broadly based on host range and specificity, DNA restriction profiles, DNA sequences from various regions of the genome, and predicted protein sequence similarities.  More recently, species demarcation criteria for alpha- and betabaculoviruses have been proposed that rely upon pairwise nucleotide distances estimated with the Kimura-2-parameter substitution model from partial sequences of three conserved baculovirus genes: late expression factor-8 (lef-8) and late expression factor-9 (lef-9), which encode subunits of the baculovirus RNA polymerase, and polyhedrin/granulin (polh/gran), which encodes the viral occlusion body matrix protein (Jehle et al., 2006b). If nucleotide distances between two viruses at these loci are less than 0.015 substitutions/site, the two baculoviruses are considered to belong to the same species. If nucleotide distances between two viruses are greater than 0.05 substitutions/site, the viruses are considered to belong to different species. If the nucleotide distances lie between 0.015 and 0.050 substitutions/site, additional characteristics of the two viruses (i.e. host range) must be considered to make a decision about their taxonomic status. The proposed criteria were originally based on an alignment of sequences from 117 separate baculovirus isolates and the phylogeny inferred from this alignment. Researchers have applied this criterion to other isolates to identify many new baculovirus species and variants of currently recognized species.

Member species

Species Virus name(s) Exemplar isolate Exemplar accession number Exemplar RefSeq number Available sequence Other isolates Other isolate accession numbers Virus abbreviation Isolate abbreviation
Adoxophyes orana granulovirus Adoxophyes orana granulovirus English AF547984 NC_005038 Complete genome AdorGV
Agrotis segetum granulovirus Agrotis segetum granulovirus DA KR584663 NC_039213 Complete genome AgseGV
Artogeia rapae granulovirus Pieris rapae granulovirus Wuhan Wuhan GQ884143 NC_013797 Complete Genome PiraGV
Choristoneura fumiferana granulovirus Choristoneura occidentalis granulovirus DQ333351 NC_008168 Complete genome ChocGV
Clostera anachoreta granulovirus Clostera anachoreta granulovirus HBHN HQ116624 NC_015398 Complete genome ClanGV
Clostera anastomosis granulovirus A Clostera anastomosis granulovirus A Henan KC179784 NC_022646 Complete genome ClasGV-A
Clostera anastomosis granulovirus B Clostera anastomosis granulovirus B KR091910 NC_038371 Complete genome ClasGV-B
Cnaphalocrocis medinalis granulovirus Cnaphalocrocis medinalis granulovirus Enping KU593505 NC_029304 Complete genome CnmeGV
Cryptophlebia leucotreta granulovirus Cryptophlebia leucotreta granulovirus CV3 AY229987 NC_005068 Complete genome CrleGV
Cydia pomonella granulovirus Cydia pomonella granulovirus Mexican 1 U53466 NC_002816 Complete genome CpGV
Diatraea saccharalis granulovirus Diatraea saccharalis granulovirus Parana-2009 KP296186 NC_028491 Complete genome DisaGV
Epinotia aporema granulovirus Epinotia aporema granulovirus JN408834 NC_018875 Complete genome EpapGV
Erinnyis ello granulovirus Erinnyis ello granulovirus S86 KJ406702 NC_025257 Complete genome ErelGV
Harrisina brillians granulovirus Harrisina brillians granulovirus M2 AF142425 NC_038372 Partial genome HbGV
Helicoverpa armigera granulovirus Helicoverpa armigera granulovirus EU255577 NC_010240 Complete genome HearGV
Lacanobia oleracea granulovirus Lacanobia oleracea granulovirus Scottish-S1 Y08294 NC_038868 Partial genome LaolGV
Mocis latipes granulovirus Mocis latipes granulovirus SouthernBrazil KR011718 NC_029996 Complete genome MolaGV-SouthernBrazil
Mythimna unipuncta granulovirus A Pseudaletia unipuncta granulovirus Hawaiian EU678671 NC_013772 Complete genome PsunGV
Mythimna unipuncta granulovirus B Mythimna unipuncta granulovirus 8 KX855660 NC_033780 Complete genome MyunGV
Phthorimaea operculella granulovirus Phthorimaea operculella granulovirus T AF499596 NC_004062 Complete genome PhopGV
Plodia interpunctella granulovirus Plodia interpunctella granulovirus Cambridge KX151395 NC_032225 Complete genome PiGV
Plutella xylostella granulovirus Plutella xylostella granulovirus K1 AF270937 NC_002593 Complete genome PlxyGV
Spodoptera frugiperda granulovirus Spodoptera frugiperda granulovirus VG008 KM371112 NC_026511 Complete genome SpfrGV
Spodoptera litura granulovirus Spodoptera litura granulovirus K1 DQ288858 NC_009503 Complete genome SpltGV
Trichoplusia ni granulovirus Trichoplusia ni granulovirus LBIV-12 KU752557 NC038375 Complete genome TnGV
Xestia c-nigrum granulovirus Xestia c-nigrum granulovirus alpha4 AF162221 NC_002331 Complete genome XecnGV
Virus names, the choice of exemplar isolates, and virus abbreviations, are not official ICTV designations.


Genus: Deltabaculovirus

Distinguishing features

Only a single species (Culex nigripalpus nucleopolyhedrovirus) has been classified in this genus.  Replication of Culex nigripalpus nucleopolyhedrovirus (CuniNPV) is restricted to host midgut epithelium, primarily in larval stages but rarely in adults. Two virion phenotypes may be characteristic of a virus species. Virions of the ODV phenotype are embedded within an occlusion body composed of a crystalline matrix of a single viral protein with no homology to polyhedrin or granulin proteins of other baculovirus genera (Perera et al., 2006, Moser et al., 2001). Each occlusion body ranges in size from 0.5 to 5 µm and contains few (1–4) or many (>50) singly-enveloped virions depending on the strain of virus, lacks the polyhedron envelope of other baculoviruses, and matures within nuclei of infected cells (Figure 1.Deltabaculovirus).

Figure 1.Deltabaculovirus. Transmission electron micrographs of occlusion bodies of Culex nigripalpus nucleopolyhedrovirus.  Scale bars: (A) 0.5 µm; (B) 0.2 µm.

Virion 

See discussion under family description.

Genome organization and replication

The CuniNPV genome is 108,252 bp and encodes at least 109 putative proteins, some of which have sequence homology with those from other baculoviruses (Afonso et al., 2001). Homologous proteins are involved in early and late gene expression, DNA replication, as well as structural and auxiliary functions. Gene orientation, order, and content in the genome of CuniNPV is different from the members of other baculovirus genera. 

Biology

Transmission of CuniNPV to larval mosquitoes is strongly influenced by divalent cations: Mg2+ is a potent enhancer of transmission whereas Ca2+ is a strong inhibitor (Becnel et al., 2001). Hosts include at least three genera of mosquitoes, but other mosquito genera and families of Diptera are likely hosts.

Species demarcation criteria

Only a single species has been classified for this genus, although other isolates that likely represent new species have been reported (Becnel and White 2007).  It is possible that criteria based on alignments of conserved gene sequences can be developed (de Araujo Coutinho et al., 2012). 

Member species

Species Virus name(s) Exemplar isolate Exemplar accession number Exemplar RefSeq number Available sequence Other isolates Other isolate accession numbers Virus abbreviation Isolate abbreviation
Culex nigripalpus nucleopolyhedrovirus Culex nigripalpus nucleopolyhedrovirus Florida1997 AF403738 NC_003084 Complete genome CuniNPV
Virus names, the choice of exemplar isolates, and virus abbreviations, are not official ICTV designations.

Related, unclassified viruses

Virus name

Accession number

Viurs abbreviation

Aedes sollicitans nucleopolyhedrovirus

JQ582836, JQ582837

AesoNPV

Uranotaenia sapphrinia nucleopolyhedrovirus

JQ582838, JQ582839

UrsaNPV

Virus names and virus abbreviations, are not official ICTV designations.

Genus: Gammabaculovirus

Distinguishing features

The nucleocapsids are enveloped singly, and multiple virions are assembled into each OB (Figure 1.Gammabaculovirus.). The virus is restricted to the host midgut and causes what was previously described in the literature as “infectious diarrhea”. Genome sequencing analyses of three viruses - Neodiprion lecontei nucleopolyhedrovirus (NeleNPV), Neodiprion sertifer nucleopolyhedrovirus (NeseNPV), and Neodiprion abietis nucleopolyhedrovirus (NeabNPV) - revealed that these viruses do not encode typical envelope fusion proteins found in other baculoviruses (Arif et al., 2011). This observation has raised the question of whether the budded virus phenotype plays a role in gammabaculovirus biology.

Figure 1.Gammabaculovirus. Transmission electron micrograph of occlusion bodies of Neodiprion abietis nucleopolyhedrovirus (micrograph courtesy of C.J. Lucarotti). Scale bar: 0.5 µm.

Virion 

See discussion under family description.

Genome organization and replication

In comparison to other baculoviruses, the genomes of members of the genus Gammabaculovirus are relatively low in G+C content (about 33%). The genomes of the three sequenced gammabaculoviruses are collinear except for a large non-syntenic region between the DNA polymerase and polyhedrin genes. This region contains genes and ORFs not shared among the three characterized genomes.

Biology

Gammabaculoviruses of species defined to date infect sawfly larvae of genus Neodiprion, order Hymenoptera, but sawflies of other genera (e.g. Gilpinia) may also contain gammabaculoviruses.

Species demarcation criteria

The two species in this genus are distinguished on the basis of differences in host range and specificity, genome sequence, gene content and gene order.  It is unclear if the nucleotide distance-based criteria developed for alpha- and betabaculoviruses can be applied to gammabaculoviruses.

Member species

Species Virus name(s) Exemplar isolate Exemplar accession number Exemplar RefSeq number Available sequence Other isolates Other isolate accession numbers Virus abbreviation Isolate abbreviation
Neodiprion lecontei nucleopolyhedrovirus Neodiprion lecontei nucleopolyhedrovirus AY349019 NC_005906 Complete genome NeleNPV
Neodiprion sertifer nucleopolyhedrovirus Neodiprion sertifer nucleopolyhedrovirus AY430810 NC_005905 Complete genome NeseNPV
Virus names, the choice of exemplar isolates, and virus abbreviations, are not official ICTV designations.

Related, unclassified viruses

Virus name

Accession number

Virus abbreviation

Gilpinia hercyniae nucleopolyhedrovirus

AY449779, AY449800

GiheNPV

Neodiprion abietis nucleopolyhedrovirus

DQ317692

NeabNPV

Virus names and virus abbreviations, are not official ICTV designations.

Authors: Baculoviridae

Robert L. Harrison*
Baculoviridae Study Group Chair
Invasive Insect Biocontrol and Behavior Laboratory
Beltsville Agricultural Research Center
USDA Agricultural Research Service
Beltsville
MD 20705
USA
Tel: 301-504-5249
E-mail: Robert.L.Harrison@ars.usda.gov

Elisabeth A. Herniou
Institut de Recherche sur la Biologie de l’Insecte
CNRS UMR 7261
Université François Rabelais
Tours 37200
France
Tel: +33 247 367381
E-mail: elisabeth.herniou@univ-tours.fr

Johannes A. Jehle
Julius Kühn Institute
Federal Research Centre for Cultivated Plants
Institute for Biological Control
Darmstadt 64287
Germany
Tel: +49 (0)6151 407 220
E-mail: johannes.jehle@julius-kuehn.de

David A. Theilmann
Summerland Research and Development Centre
Agriculture and Agri-Food Canada
Summerland
BC V0H 1Z0
Canada
Tel: 250-494-6395
E-mail: david.theilmann@agr.gc.ca

John P. Burand
Department of Microbiology
University of Massachusetts-Amherst
Amherst
MA 01003
USA
Tel: 413-545-3629
E-mail: jburand@microbio.umass.edu

James J. Becnel
Center for Medical, Agricultural and Veterinary Entomology
USDA Agricultural Research Service
Gainesville
FL 32608
USA
Tel: 352-374-5961
E-mail: James.Becnel@ars.usda.gov

Peter J. Krell
Department of Molecular and Cellular Biology
University of Guelph
Guelph
Ontario N1G 2W1
Canada
Tel: 519-824-4120 x53368 or 53264
E-mail: pkrell@uoguelph.ca

Monique M. van Oers
Laboratory of Virology
Wageningen University
Wageningen 6709 PD
The Netherlands
Tel: +31317485082
E-mail: monique.vanoers@wur.nl

Joseph D. Mowery
Electron and Confocal Microscopy Unit
Beltsville Agricultural Research Center
USDA Agricultural Research Service
Beltsville
MD 20705
USA
Tel: 301-504-9027
E-mail: Joseph.mowery@ars.usda.gov

Gary R. Bauchan
Electron and Confocal Microscopy Unit
Beltsville Agricultural Research Center
USDA Agricultural Research Service
Beltsville
MD 20705
USA
Tel: 301-504-6649
E-mail: gary.bauchan@ars.usda.gov

* To whom correspondence should be addressed

The chapter in the Ninth ICTV Report, which served as the template for this chapter, was contributed by Herniou, E.A., Arif, B.M., Becnel, J.J., Blissard, G.W., Bonning, B., Harrison, R., Jehle, J.A., Theilmann, D.A. and Vlak, J.M.


Resources: Baculoviridae

Websites:

Baculovirus Molecular Biology, 3rd edition: https://www.ncbi.nlm.nih.gov/books/NBK114593/

Sequence alignments and tree files:

Figure 3.Baculoviridae:

Tree file (newick format)

Alignment file (FASTA format)

Individual gene alignment pre-concatenation (ZIP)


Further reading: Baculoviridae

Braunagel, S. C. & Summers, M. D. (2007). Molecular biology of the baculovirus occlusion-derived virus envelope. Curr Drug Targets 8, 1084-1095. [PubMed]

Herniou, E. A. & Jehle, J. A. (2007). Baculovirus phylogeny and evolution. Curr Drug Targets 8, 1043-1050. [PubMed]

Herniou, E. A., Olszewski, J. A., Cory, J. S. & O'Reilly, D. R. (2003). The genome sequence and evolution of baculoviruses. Annu Rev Entomol 48, 211-234. [PubMed]

Miller, L. K. (1997). The Baculoviruses. New York: Plenum Press.

van Oers, M. M. & Vlak, J. M. (2007). Baculovirus genomics. Curr Drug Targets 8, 1051-1068. [PubMed]


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Citation: Baculoviridae

A summary of this ICTV Report chapter has been published as an ICTV Virus Taxonomy Profile article in the Journal of General Virology, and should be cited when referencing this online chapter as follows:

Harrison, R.L., Herniou, E.A., Jehle, J.A., Theilmann, D.A., Burand, J.P., Becnel, J.J., Krell, P.J., M. van Oers, M., Mowery, J.D., Bauchan, G.R., and ICTV Report Consortium. 2019, ICTV Virus Taxonomy Profile: BaculoviridaeJournal of General Virology, 99: 11851186.