Iridoviruses

Introduction | Taxonomy | Morphology | Genome | Replication | Gene expression | Pathogenesis

Introduction
The family Iridoviridae contains a diverse array of large icosahedral viruses that replicate in the cytoplasm of infected cells. The word Iridoviridae is derived from Iris who was the Greek goddess of the rain ow. This is due to the !rain ow like! iridescence o served in heavily infected insects and pelleted samples of inverte rate iridoviruses.

Larvae of the grass grub Costelytra zealandica displaying blue colouration of the hindgut due to iridovirus infection.

Pellet of purified Tipula iridescent virus This iridescence facilitated the first detection of an iridovirus when "laude Rivers# in March of $%&'# discovered crane fly larvae (Tipula spp) glowing with patches of lue colouration. Iridoviruses have since een isolated from oth inverte rate and non*mammalian verte rate

*$) 2rog .1) The current I"T. + common feature of most of these hosts is the a. N nucleus! "# viroplasmic centre Ta$onomy The family Iridoviridae is comprised of four genera# two infecting verte rates and two infecting inverte rates. The wide host range displayed y mem ers of this family has resulted in isolates of a single virus species eing descri ed and named more than once ecause of their isolation from different hosts.uacultural practices and ecause of their potential use in the iological control of insect pests.1) 0ymphocystivirus type $ (0"3.irus 1 (2. Paracrystalline array of virus particles within infected cell. Sf21 tissue culture cell showing cytoplasmic localisation of assembling Wiseana iridescent virus particles. Iridoviruses have received attention ecause of the pro lems they pose to a. This array gives rise to the iridescent phenomenon.hosts.uatic or moist environment in which they are found. %enus Iridovirus Chloriridovirus Lymphocystivirus Ranavirus "ernacular name -mall iridescent insect virus 0arge iridescent insect viruses 0ymphocystis disease virus 2rog virus &ost species Inverte rates (mainly insects) Mos./) Mos.uito iridescent virus (I. list of recognised Iridoviridae mem ers contains much redundant data.uitos 2ish +mphi ia Type species Chilo iridescent virus (I. The host of isolation also confuses taxonomy within the family with recently identified .

homology to the #-I" protein to white for +.fish iridoviruses seemingly more related to ranaviruses than to lymphocystiviruses. homology.ues. )ars represent the ma(or capsid proteins from each virus with greyscales ranging from blac* for 1++. . 2or example# the P"R amplification and se.uencing of a &55 p fragment of the ma4or capsid protein gene from a num er of inverte rate iridoviruses suggests the genus Iridovirus can e further divided into three groups. 'epresentation of the homologies between invertebrate iridovirus ma(or capsid genes highlighting the region used for phylogenetic analysis. The pitfalls in iridovirus taxonomy are only 4ust eginning to e addressed using molecular techni.

(I. (I.7' (+I.$) -I. (I.15 I.7) "I. (I.) Iridovirus I.%) I.ealand 9ales -outh +frica India >-+ >-+ >-+ "<I.ealand :ew .1$ Iridovirus Iridovirus Iridovirus . (I.#P se/uence #ountries of virus isolation "irus TI. (I.$/) Iridovirus = I.77 %enus Iridovirus Iridovirus Iridovirus Iridovirus Iridovirus &ost crane fly scare gru rice stem orer porina caterpillar grass gru lack fly lack eetle honey ee meal worm corn earworm woodlice #ountry of isolation 6ngland +ustralia 8apan :ew .Phylogenetic tree of the genus Iridovirus based on partial .7% I./) 9I.71) Iridovirus I.

suggested that a structure represented y $'@7 su units may e more appropriate.uired y udding through host mem ranes. 0arger iridoviruses reak down into larger ut morphologically similar su units.elvet ean caterpillar +rgentina 8apanese eetle +<ores 9orld wide Lymphocystivirus flounder . P4I.*$ Iridovirus Iridovirus .1. + common feature of all iridoviruses is the presence of a ma4or capsid protein of around &5 k3a that accounts for up to '&A of total virion protein.# and "I.orphology 0lectron micrograph of a typical iridovirus Iridoviruses are large ($75 to 155 nm in diameter) non*occluded viruses with icosahedral symmetry. These o servations led 9rigley to propose the following $&/7 morphological su unit structure for -I. Prolonged storage of Sericesthis iridescent virus (-I. 0"3.*$# MI.. 2i rillar structures have also een o served protruding from capsid su units of 0"3. .) in distilled water at 'B" led to the disintegration of virions into triangular# pentagonal and linear fragments consisting of &&# $/# and % su units respectively. -ome# ut not all# viruses possess an outer envelope ac. 0ater studies on TI.# ut not from 2. +n iridovirus virion is composed of three concentric domains? an outer proteinaceous capsid# an intermediate lipid mem rane with associated polypeptides# and a central core containing 3:+*protein complexes. 3epending on the detection method used iridoviruses possess etween 7& to @& structural proteins ranging in molecular weight from $7#555 to $&5#555 k3a.+gI.

irol. &C$71*$1') %enome Iridoviruses contain a single copy linear ds3:+ genome that ranges in si<e from $&5 to 7D5 k p depending on viral species.ue within the eucaryotic viruses in that they are terminally redundant and cyclically permuted. .Schematic diagram adapted from N. The genomes appear uni.%. . This structure is a result of the resolution of genome concatamers during 3:+ replication (see replication). Gen. 1rigley 1232 (8.

and 0"3. This genomic structure has een found in all iridoviruses so far studied.uenced and all were predicted to form hairpin structures. -u tle variations do occur etween different genera in the cyclic permutation. The resolution of this concatamer (") results in packaged 3:+ lengths that contain a complete genome as well as duplicated copies of some genes (terminal redundancy).) appear uni. The ends of each of these packaged 3:+s differs from one virus particle to the next (cyclic permutation).ue se.4 simplistic view of terminal redundancy and cyclic permutation. The packaged 3:+ ends within a population of 2.uences or what its role is. Plasmid rescue experiments identified six origins of replication in "I. +lthough packaging occurs in the cytoplasm of infected cells# a nuclear stage is also present.ue in that they contain at least two genomic fragments# one of genome length and another of approximately $$ k p. The particles of Tipula iridescent virus (TI. . 'eplication The replication of iridoviruses comes mainly from studies of 2.1 and this has ecome the model for iridovirus replication.*$ the permutation is completely random covering $55A. Three of these origins have een se.# a feature consistent with cyclic permutation. This smaller su genomic fragment hy ridises to defined regions of the genome ut it is not known whether it also contains uni. 3uring replication multiple copies of a hypothetical viral genome consisting of $5 genes (+) forms a long concatamer (=).1 are confined to regions representing 7&A of the genome whereas in "I.

%oorha 1272 (8. 1) Parental 3:+ is used to produce genome and greater than genome length 3:+. &) "oncatamers are resolved into packaged lengths# possi ly y a headful packaging approach.5"6 replication model adapted from '.iral 3:+ is transported to the cell nucleus where host macromolecular synthesis is rapidly shutdown. . ') Progeny 3:+ is transported into the cytoplasm where large concatamers of viral 3:+ are formed y recom ination. This ecomes the template for cytoplasmic replication. Transcription of very late transcripts may also take place in the cytoplasm.irus particles enter the cell y pinocytosis and uncoating occurs. 7) .irions exit the cell y udding or cell lysis. .irol. . Transcription is initiated y virally modified host R:+ polymerase II. '1(7)C&$%*&7D) $) .

Possible recombination dependent iridovirus replication. 7$C1'@*1@$) . These single stranded regions are capa le of recom ination within the same duplex (") or with another 3:+ molecule (=). Mosig $%D@ (+nn. Modified from G. Rev. Erigin dependent replication results in the production of duplex 3:+ with single stranded 1F ends (+). Genet. In this this way these 1F ends serve as primers for further 3:+ replication. The result is a large interlinked and replicating 3:+ concatamer.

Iridoviruses from the lackfly# Simulium spp.1 is not known to cause disease in naturally occurring frog populations although frog em ryos and larvae die within $& days if inoculated with virus. 5rog virus 6 transcription control model Pathogenesis 0ittle is known a out the pathogenesis of iridoviruses. In a lethal infection y insect iridoviruses the fat odies and haemocytes are the initial sites of replication# this leading to a systemic infection. 2. Insects ecome flaccid and iridescent @*$5 days post*infection although death may take 1 weeks or longer.%ene e$pression -tudies of oth verte rate and inverte rate iridoviruses show transcription occurs in a temporal manner with three stages? immediate*early# delayed*early# and late. . There is oth positive induction and some negative feed ack on transcription at each level y translational products of other temporal stages.# are found in two states# covert (inapparent) and patent (lethal)# the ratios of each dependent on environmental conditions and host densities. 2.1 can cause edema of tadpole tails ut has no apparent effect if inoculated into adult frogs. The pathogenesis is# however# temperature dependent and iridoviruses are thus confined to poikilothermic hosts.

.*$ causes a nonlethal viral disease of many marine and freshwater fish.*$ appear to e non*lethal# high mortality rates amongst various frog and fish populations have een caused y other viruses of the Iridoviridae family. +lthough 2.0"3. Infected fish often have nonmalignant tumour*like growths and rasp erry*like lesions associated with their skin.1 and 0"3. + characteristic of the disease is the presence of enormously enlarged host cells (cells can increase in volume y a factor of $5/).