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Viruses are obligate intracellular parasites. They consist of a DNA or RNA genome surrounded by protein. They may also have an outer-membrane lipoprotein envelope. Viruses can replicate only within cells because their nucleic acid does not encode many enzymes necessary for the metabolism of proteins, carbohydrates, or lipids or for the generation of high-energy phosphates. Typically, viral nucleic acids encode messenger RNA (mRNA) and proteins necessary for replicating, packaging, and releasing progeny virus from infected cells.

Viruses differ from virusoids, viroids, and prions. Virusoids are nucleic acids that depend on cells and helper viruses for packaging their nucleic acids into virus-like particles. Viroids are naked, cyclical, mostly double-strand small RNAs that appear to be restricted to plants, spread from cell to cell, and are replicated by cellular RNA polymerase II. Prions (Chap. 109) are abnormal proteins that propagate and cause disease by altering the structure of a normal cell protein. Prions cause neurodegenerative diseases such as Creutzfeldt-Jakob disease, Gerstmann-Straüssler disease, kuru, and human or bovine spongiform encephalopathy (“mad cow disease”).


Viral genomes may consist of single- or double-strand DNA, single- or double-strand RNA, single-strand or segmented antisense RNA, or double-strand segmented RNA. Viral nucleic acids may encode only a few genes or more than 100. Sense-strand viral RNA genomes can be translated directly into protein, whereas antisense RNAs must be copied into translatable RNA. Sense and antisense genomes are also referred to as positive-strand and negative-strand genomes, respectively. Viral nucleic acid is usually associated with virus-encoded nucleoprotein(s) in the virus core. Viral nucleic acids and nucleoproteins are almost always enclosed in a protein capsid. Because of the limited genetic complexity of viruses, their capsids are usually composed of multimers of identical capsomeres made up of one or a few proteins. Capsids have icosahedral or helical symmetry. Icosahedral capsid structures approximate spheres and have two-, three-, or fivefold axes of symmetry, whereas helical capsid structures have only a twofold axis of symmetry. The nucleic acid, nucleoprotein(s), and protein capsid together are called a nucleocapsid.

Many viruses are composed of a nucleic acid core and a capsid. For these viruses, the outer capsid surface mediates contact with uninfected cells’ plasma membranes. Other viruses are more complex and have an outer phospholipid, cholesterol, glycoprotein, and glycolipid envelope that is derived from virus-modified infected cell membranes. Cell nuclear, endoplasmic reticulum, Golgi, or plasma membranes that become parts of the viral envelope have usually been modified during infection by the insertion of virus-encoded glycoproteins, which mediate contact of enveloped virus with uninfected cell surfaces. Matrix or tegument proteins may fill the space between the nucleocapsid and the outer envelope of the virus.

Enveloped viruses are usually sensitive to lipid solvents or detergents that can dissolve the envelope, whereas viruses with protein nucleocapsid exteriors may be somewhat detergent resistant. A schematic ...

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