Polydnaviridae

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Polydnaviruses
GiBV-em.jpg

Polydnaviruses

Systematics
Classification : Viruses
Area : unclassified
Empire : unclassified
Phylum : unclassified
Order : unclassified
Family : Polydnaviridae
Taxonomic characteristics
Genome : dsDNA
Baltimore : Group 1
Scientific name
Polydnaviridae
Left

The polydnaviruses are a family of enveloped viruses with a double-stranded, segmented DNA genome , which are integrated into the genome of some parasitic wasps and serve as a viral vector to influence the hosts of the parasitic wasp larvae. The family contains two genera with little homology to one another, the Ichnoviruses with a single lipid bilayer and the Bracoviruses with a double lipid bilayer in their respective virus envelope. The former occur in Ichneumonidae , while the latter occur in Braconidae .

structure

While Ichnoviruses have an egg-shaped shape in the transmission electron microscope ( Ascoviridae -like morphology ), Bracoviruses are rod-shaped ( Baculoviridae -like). The emergence of the polydnaviruses is dated between 64 and 100 million years before our era and is now only transmitted to the offspring (vertical infection ). The viral genome is integrated into the genome of the parasitic wasp, while the polydnavirus virions are only formed in the calyx cells of the ovaries and are injected into the parasitized caterpillar together with the wasp eggs. The virion contains multiple copies of viral dsDNA in segmented, circular and supercoiled form with lengths between 2 Kbp and 31 Kbp. Depending on the species, one to several capsids occur per virion, i.e. within a virus envelope. Bracoviruses leave the calyx cells by lysis , while Ichnoviruses leave the calyx cells by budding .

Parasitism and symbiosis

Polydnavirus injection

Polydnaviruses are part of a unique host-pathogen relationship between parasitoid wasps, the parasitized caterpillars (mostly Lepidoptera ) and the symbiotic virus of the ichneumon fly. The caterpillars are hosts of the parasitoid parasitic wasp larvae, while the parasitic wasps are symbiotic hosts for the polydnaviruses. At the same time, polydnaviruses are pathogens for the caterpillars, which is the symbiotic benefit for the parasitic wasps. The polydnavirus virions are only formed in the calyx cells of the ovaries and are injected into the parasitized caterpillar together with the wasp eggs. The virion does not contain all the necessary genes for replication , but does contain various virulence factors derived from the parasitic wasp genome , including shortened versions with homology to caterpillar genes that inhibit their function . The virus modulates the immune system , metabolism and behavior of the parasitized caterpillar without being replicated in it. Polydnaviruses and parasitic wasps thus have a symbiotic mutualism , because the virus serves as a vector for transient gene transfer , while the wasps only reproduce in their ovaries.

Inhibition of defense reactions

The modulation of the immune response to the injected eggs in the parasitized hosts prevents the defense reaction that usually occurs. This consists of an encapsulation of the eggs with the hematocytes of the mixocoel , which then reduces the oxygen supply to the foreign body through further hematocytes via the phenol oxidase with melanin , whereby the development of a pathogen is slowed down or stopped. Polydnaviruses can, among other things, inactivate or destroy NF-κB in hematocytes, which weakens this defense. The polydnavirus from Cotesia rubecula encodes the protein CrV1 , which depolymerizes actin filaments in hematocytes , which limits their mobility and stability. The Bracovirus of the Microplitis demolitor (MdBV) induces apoptosis in hematocytes by means of PTP-H2 , which results in less defense and more liquid food is available. Furthermore, the phagocytosis and adhesion of the haematocytes is reduced by the gene GLC1.8 , which further reduces the resistance . By Egf1.0 the phenol oxidase is inhibited, which inhibits the formation of melanin in the course of the immune defense.

The Ankyris protein inhibits the effects of antiviral peptides . In some Ichnoviruses, Vankyrin inhibits the induction of apoptosis. The viral innexin-related protein Vinnexin changes the communication between the cells of the caterpillar at the gap junctions . In addition, polydnaviruses induce hyperglycemia in the host , which improves the food supply for the parasitic wasp larvae.

Furthermore, polydnaviruses lead to behavioral changes in the parasitized caterpillars, which after infection refrain from migrating and pupating in order to protect the parasitic wasp larvae that have hatched and pupate from predators or other parasitoids until their own energy reserves are used up and the caterpillar dies. Polydnaviruses are the first known case of a gene transfer vector that is used by parasitoids to modify the host's behavior in addition to modulating the immune system and metabolism . It is also the first known case that hosts protect the offspring of their parasites or parasitoids outside of their bodies until they give up themselves.

Virus-like particles

Some polydnaviruses only produce virus-like particles (VLP) without any nucleic acids they contain , which in themselves dampen the immune response, e.g. B. in Venturia canescens ( Ichneumonidea ) and Leptopilina sp. ( Figitidaea ). The VLP of the canescens vein (VcVLP1, VcVLP2, VcNEP…) are produced in calyx cells in order to be released into the fallopian tube. Since the DNA sequences of the VLP proteins show more similarities with Hymenoptera proteins than with viral proteins, a non-viral origin has been assumed.

The wasp Leptopilina heterotoma secretes VLP, which can penetrate the lamellocytes.

Occurrence

Polydnaviruses occur in various subfamilies of the parasitic wasps, including

  • Bracovirus in:
  • Ichnovirus in:

Systematics

Internal system

As of February 2019, the International Committee on Taxonomy of Viruses (ICTV) reports that the Polydnaviridae family consists of the following two genera:

  • Family Polydnaviridae

External system

Nucleic acid analyzes suggest a very long association between viruses and wasps (more than 70 million years).

With regard to the co-evolution of wasps and viruses, two contradicting proposals have been made.

  • The first suggestion is based on the assumption that the virus is derived from wasp genes. Many non-polydnavirus parasitoid wasps inject proteins with many equivalent functions; H. suppress the immune response of the host to the Parasitenei and therefore English virus-like particles are (VLPs) called ( see below ). In this model, the braconid and ichneumonid wasps have packed genes for these functions into the viruses - which essentially leads to a gene transfer system in which the caterpillar itself produces the immunosuppressive factors. In this scenario, the structural proteins of the viruses ( capsids ) were probably "borrowed" from existing viruses.
  • The alternative proposal assumes that wasp ancestors developed a beneficial symbiosis with an existing virus that eventually resulted in the virus being integrated into the wasp's genome . The genes responsible for virus replication and the capsids were then unnecessary and at some point no longer included in the virus genome.

This second hypothesis is supported by the different morphologies between IV and BV, which suggests that the two genera have different origins. The IV bears remarkable similarities to, and is believed to have evolved from, ascoviruses , while the BV may have evolved from the Nudiviridae about 100 million years ago . The Polydnaviridae (i.e. the group of Bracoviruses) appear according to Koonin et al. (2015 and 2019) together with the Nimaviridae , Hytrosaviridae , Baculoviridae and the Nudiviridae to form an as yet unnamed family group, for which the authors propose the following family tree:



Nimaviridae


   

Hytrosaviridae


   

Baculoviridae


   

Nudiviridae


   

Polydnaviridae : Bracoviruses




Template: Klade / Maintenance / 3

Template: Klade / Maintenance / Style

The two genera of viruses in the family are actually not phylogenetically related, which means that this taxon may need some overhaul.

Web links

Individual evidence

  1. ICTV: ICTV Master Species List 2019.v1 , New MSL including all taxa updates since the 2018b release, March 2020 (MSL # 35)
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  30. A. Bézier, M. Annaheim, J. Herbinière, C. Wetterwald, G. Gyapay, S. Bernard-Samain, P. Wincker, I. Roditi, M. Heller, M. Belghazi, R. Pfister-Wilhem, G Periquet, C. Dupuy, E. Huguet, AN Volkoff, B. Lanzrein, JM Drezen: Polydnaviruses of braconid wasps derive from an ancestral nudivirus. In: Science (2009), Vol. 323 (5916), pp. 926-30, PMID 19213916 , doi: 10.1126 / science.1166788 .
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