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lysis breakage that can be replicated to produce progeny genomes that will be packaged into newly produced virions. Replication details vary among the different types of viruses.

The ss positive-sense DNA of parvoviruses is copied by host DNA polymerase (the enzyme that replicates DNA) in the nucleus into a negativesense DNA strand. This in turn serves as a template for mRNA and progeny DNA synthesis. The genomes of larger DNA viruses, with the exception of the poxviruses, are also transcribed and replicated in the nucleus by a combination of viral and host enzymes (for example, DNA-dependent RNA polymerses for transcription of mRNAs, DNA-dependent DNA polymerase for genome replication).

Positive-polarity RNA virus genomes can be translated directly, but for effective progeny production additional rounds of RNA replication via a negative-stranded intermediate are required. This is accomplished by a viral transcriptase (RNA-dependent RNA polymerase) and associated cofactors. Single-stranded negative-sense RNA viruses of animals must also carry a viral transcriptase to transcribe functional mRNAs and subsequently produce proteins, since this RNA-to-RNA enzymatic activity is typically lacking in animal cells.

Retroviruses are unique among viruses in that the genome is diploid, meaning that two copies of the positive-polarity RNA genome are in each virus particle. The genomic RNA is not translated into protein, but rather serves as a template for reverse transcription, which produces a double-stranded DNA via a viral reverse transcription enzyme. The DNA is subsequently integrated into the host cell chromosomal DNA. Hepadnaviruses also encode a reverse transcriptase, but replication occurs inside the virus particle producing the particle-associated genomic DNA.

Assembly, Maturation, and Release. As viral proteins and nucleic acids accumulate in the cell, they begin a process of self-assembly. Viral self-assembly was first demonstrated in a seminal series of experiments in 1955, wherein infectious particles of tobacco mosaic virus spontaneously formed when purified coat protein and genomic RNA were mixed. Likewise, poliovirus capsomers are known to self-assemble to form a procap-sid in the cytoplasm. Progeny positive-strand poliovirus RNAs then enter this nascent particle. "Chaperone" proteins (chaparonins) of the cell play a critical role in facilitating the assembly of some viruses. Their normal role is to help fold cellular proteins after synthesis.

The maturation and release stages of the replication cycle may occur simultaneously with the previous step, or may follow in either order. Many viruses assemble their various components into "immature" particles. Further intracellular or extracellular processing is required to produce a mature infectious particle. This may involve cleavage of precursors to the structural proteins, as in the case of retroviruses.

Viruses that are not enveloped usually depend upon disintegration or lysis of the cell for release. Enveloped viruses can be released from the cell by the process of budding. In this process the viral capsid and usually a matrix layer are directed to a modified patch of cellular membrane. Interactions between the matrix proteins and/or envelope proteins drive envelopment. In the case of viruses that bud at the cell surface, such as some

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