Nudivirus

In 2007, the genus Nudivirus was proposed to include viruses similar to the Oryctes rhinoceros virus.[2] The nudiviruses were classified as the family Nudiviridae in 2013.[3]

All 3 sequenced nudivirus have 33 open reading frames in common. Gene content comparaison and phylogenetic analyses show that Nudivirus share 20 cores genes with baculovirus and form a monophyletic sister group with them. Fossil calibration estimate this association arose 100 million years ago(Mya), while the last common ancestor of BVs, nudivirus, and baculovirus existed approximately 312 Mya. Baculoviruses and nudiviruses differ in gene content, genome organization, cytopathology, infection of adults and most likely in host range.[4] The 20 cores genes common in both Baculovirus and Nudivirus are involved in RNA transcription, DNA replication, virion structural components and many other functions.[5] Gene content and sequence similarity suggest that the nudiviruses GbNV, HzNV-1, and OrNV form a monophyletic group of nonoccluded double-stranded DNA viruses, which separated from the baculovirus lineage before this radiated into dipteran-, hymenopteran-, and lepidopteran-specific clades of occluded nucleopolyhedroviruses and granuloviruses.[4]

This genus is characterized by rod-shaped and enveloped nucleocapsids and its members replicate in the nucleus of infected host cells. In some parasitoid wasp species, a nudivirus genome, in proviral form, is integrated the wasp genome and produces virus like particles called polydnaviruses that are injected into lepidopteran larvae and are thought to facilitate parasitization of the larvae (see below). Nudivirus are known to infect preferentially insects and marine arthropods.[2] Transmission of nudiviruses occurs generally by feeding and/or mating. Infections can be lethal for the larvae and can possibly reduce the fitness of the host by reducing offspring production and survival among adults.[6]

• Drosophila innubila nudivirus – Drosophila innubila (Diptera)
• Gryllus bimaculatus nudivirus – black cricket (Teleogryllus commodus)
• Helicoverpa zea nudivirus 1 – cotton bollworm
• Helicoverpa zea nudivirus 2 – cotton bollworm
• Homarus gammarus nudivirus - common lobster (Homarus gammarus)
• Oryctes rhinoceros nudivirus – rhinoceros beetle (Dynastinae species)
• Penaeus monodon nudivirus – a shrimp
• Dikerogammarus haemobaphes nudivirus – an amphipod crustacean (Dikerogammarus haemobaphes)[7]

In many organisms, apoptosis can be regarded as an early cellular defense mechanism against a strong viral infection. Although, Heliothis zea Nudivirus 1 (HzNV-1 or Hz-1 virus), a relatively broad host range nudivirus, has been shown to block an induced-apoptosis gene (hhi1), and so, prevent his own death. A functional antiapoptosis gene, (Hz-iap2), has been highlighted to suppress the hhi1 gene which can either activate the expression of prodeath gene or alter the cell cycle within the host, resulting in the death of the cell. A second inhibitor gene (Ac-iap2) to the hhi1 gene has been also discovered, Ac-iap2 was speculated to be a nonfunctional iap, raising many questions regarding its functionality.[8]

The parts of the cycle that relate to the transmission of wasp genomic DNA and therefore to proviral DNA are shown by the claret arrows. Those parts of the cycle that apply only to virus replication and assembly/transmission are shown by the green arrows. Polydnavirus (PDVs) are associated to endoparasitic wasps from the Ichneumoninae family (Hymenoptera). There are two genera of PDVs: The ichnoviruses occur in ichneumonid wasps species and bracoviruses in braconid wasps. The polydnaviridae appear to replicate and accumulate in the ovarian calyx epithelial cells and calyx fluid of the wasps and do not replicate in the lepidopteran hosts of the wasps, although their gene products are expressed there. The packaged Virus is injected along with the wasp egg into the body cavity of a lepidopteran host caterpillar and infects cells of the caterpillar.[9] The infection does not lead to replication of new viruses, rather it affects the caterpillar's Immune system.

Without the virus infection, Phagocyte Hemocyte (blood cells) will encapsulate and kill the wasp egg but the immune suppression caused by the virus allows for survival of the wasp egg, leading to hatching and complete development of the immature wasp in the caterpillar. Additionally, genes expressed from the polydnavirus in the parasitised host alter host development and metabolism to be beneficial for the growth and survival of the parasitoid larva.[10] Thus the virus and wasp have a Symbiosis (mutualistic) relationship. It appears in most cases that the PDVs DNA is required for successful Parasitism of Lepidoptera by the wasps. So, the full genome of the virus is integrated into the genome of the wasp and the virus only replicates in specific cells in the female wasp's reproductive system. As a result, the polydnaviral DNA has experienced spectacular reproductive success; it is now thought to be represented in species of many ichneumonoid wasps.

The conservation of this viral machinery in different braconid wasp lineages sharing polydnaviruses suggests that parasitoid wasps incorporated a nudivirus-related genome into their own genetic material.[11]