1.4 THE VIRUS MULTIPLICATION CYCLE
We now know a great deal about the processes which occur
during the multiplication of viruses within single cells. The precise details
vary for individual viruses but have in common a series of events marking
specific phases in the multiplication cycle. These phases are summarized in
Fig. 1.3 and are considered in detail in section II of this book. The first
stage is that of attachment when the virus attaches to the potential host cell.
The interaction is specific, with the virus attachment protein(s) binding to
target receptor molecules on the surface of the cell. The initial contact
between a virus and host cell is dynamic and reversible, and often involves
weak electrostatic interactions. However, the contacts quickly become much
stronger with more stable interactions which in some cases are essentially irreversible.
The attachment phase determines the specificity of the virus for a particular
type of cell or host species. Having attached to the surface of the cell, the
virus must effect entry to be able to replicate in a process called penetration
or entry. Once inside the cell the genome of the virus must become available.
This is achieved by the loss of many, or all, of the proteins that make up the
particle in a process referred to as uncoating. For some viruses the entry and
uncoating phases are combined in a single process. Typically these first three
phases do not require the expenditure of energy in the form of ATP hydrolysis.
Having made the virus genome available it is now used in the biosynthesis phase
when genome replication, transcription of mRNA, and translation of the mRNA
into protein occur. The process of translation uses ribosomes provided by the
host cell and it is this requirement for the translation machinery, as well as
the need for molecules for biosynthesis, that makes viruses obligate
intracellular parasites. The newly synthesized genomes may then be used as
templates for further rounds of replication and as templates for transcription
of more virus mRNA in an amplification process which increases the yield of
virus from the infected cells. When the new genomes are produced they come
together with the newly synthesized virus proteins to form progeny virus
particles in a process called assembly. Finally, the particles must leave the
cell in a release phase after which they seek out new potential host cells to
begin the process again. The particles produced within the cell may require
further processing to become infectious and this maturation phase may occur before
or after release. Combining the consideration of the steps which make up a
virus multiplication cycle with the information in the graph of the results of
a single step growth curve it can be seen that during the eclipse phase the
virus is undergoing the processes of attachment, entry, uncoating, and
biosynthesis. At this time the cells contain all of the elements necessary to
produce viruses but the original infecting virus has been dismantled and no new
infectious particles have yet been produced. It is only after the assembly step
that we see virus particles inside the cell before they are released and appear
in the medium
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