%0 Journal Article %T Structural Differences Observed in Arboviruses of the Alphavirus and Flavivirus Genera %A Raquel Hernandez %A Dennis T. Brown %A Angel Paredes %J Advances in Virology %D 2014 %I Hindawi Publishing Corporation %R 10.1155/2014/259382 %X Arthropod borne viruses have developed a complex life cycle adapted to alternate between insect and vertebrate hosts. These arthropod-borne viruses belong mainly to the families Togaviridae, Flaviviridae, and Bunyaviridae. This group of viruses contains many pathogens that cause febrile, hemorrhagic, and encephalitic disease or arthritic symptoms which can be persistent. It has been appreciated for many years that these viruses were evolutionarily adapted to function in the highly divergent cellular environments of both insect and mammalian phyla. These viruses are hybrid in nature, containing viral-encoded RNA and proteins which are glycosylated by the host and encapsulate viral nucleocapsids in the context of a host-derived membrane. From a structural perspective, these virus particles are macromolecular machines adapted in design to assemble into a packaging and delivery system for the virus genome and, only when associated with the conditions appropriate for a productive infection, to disassemble and deliver the RNA cargo. It was initially assumed that the structures of the virus from both hosts were equivalent. New evidence that alphaviruses and flaviviruses can exist in more than one conformation postenvelopment will be discussed in this review. The data are limited but should refocus the field of structural biology on the metastable nature of these viruses. 1. Background 1.1. Arbovirus Evolution The arboviruses are not a taxonomic classification, but rather a grouping based on viral transmission through an insect vector to infection of a vertebrate host. The arboviruses contain members of the Togaviridae, Flaviviridae, Bunyaviridae, Rhabdoviridae, Reoviridae, and Orthomyxoviridae and are also represented by a single DNA virus, African swine fever virus family Asfarviridae of genus Asfivirus http://ictvonline.org/virusTaxonomy.asp. Evidence exists that arboviruses from the alphavirus lineage evolved from plant viruses [1, 2] which adapted to growth in insects [3]. Hematophagous insect viruses then acquired the ability to infect vertebrates, thus adapting from separate kingdoms (plant to insect) as well as phyla (insect to vertebrate) [4]. Members of the Bunyaviridae still maintain the plant to insect cycle [5¨C7] as well as the insect only cycle [8¨C10]. Arbovirus members of the flaviviruses are believed to have emerged about 1000 years ago in a nonhuman primate to mosquito cycle [11, 12] from predecessors that date at least 85,000 years [8]. It has been suggested that each of the 4 dengue serotypes (DEN1-4) adapted to humans independently only a %U http://www.hindawi.com/journals/av/2014/259382/