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A viroplasm, sometimes called "virus factory" or "virus inclusion", is an
inclusion body Inclusion bodies are aggregates of specific types of protein found in neurons, a number of tissue cells including red blood cells, bacteria, viruses, and plants. Inclusion bodies of aggregations of multiple proteins are also found in muscle cells ...
in a
cell Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery ...
where
viral replication Viral replication is the formation of biological viruses during the infection process in the target host cells. Viruses must first get into the cell before viral replication can occur. Through the generation of abundant copies of its genome an ...
and assembly occurs. They may be thought of as viral factories in the cell. There are many viroplasms in one infected cell, where they appear dense to
electron microscopy An electron microscope is a microscope that uses a beam of accelerated electrons as a source of illumination. As the wavelength of an electron can be up to 100,000 times shorter than that of visible light photons, electron microscopes have a hi ...
. Very little is understood about the mechanism of viroplasm formation.


Definition

A viroplasm is a perinuclear or a
cytoplasm In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. ...
ic large compartment where
viral replication Viral replication is the formation of biological viruses during the infection process in the target host cells. Viruses must first get into the cell before viral replication can occur. Through the generation of abundant copies of its genome an ...
and assembly occurs. The viroplasm formation is caused by the interactions between the virus and the infected
cell Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery ...
, where viral products and cell elements are confined.


Groups of viruses that form viroplasms

Viroplasms have been reported in many unrelated groups of
Eukaryotic Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
viruses that replicate in
cytoplasm In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. ...
, however, viroplasms from plant viruses have not been as studied as viroplasms from animal viruses. Viroplasms have been found in the
cauliflower mosaic virus Cauliflower mosaic virus (CaMV) is a member of the genus ''Caulimovirus'', one of the six genera in the family ''Caulimoviridae'', which are pararetroviruses that infect plants. Pararetroviruses replicate through reverse transcription just like ...
,
rotavirus ''Rotavirus'' is a genus of double-stranded RNA viruses in the family ''Reoviridae''. Rotaviruses are the most common cause of diarrhoeal disease among infants and young children. Nearly every child in the world is infected with a rotavirus ...
, vaccinia virus and the
rice dwarf virus Rice dwarf virus (RDV) is a plant pathogenic virus of the family Reoviridae ''Reoviridae'' is a family of double-stranded RNA viruses. Member viruses have a wide host range, including vertebrates, invertebrates, plants, protists and fungi. T ...
. These appear electron-dense under an
electron microscope An electron microscope is a microscope that uses a beam of accelerated electrons as a source of illumination. As the wavelength of an electron can be up to 100,000 times shorter than that of visible light photons, electron microscopes have a hi ...
and are insoluble.


Structure and formation

Viroplasms are localized in the perinuclear area or in the
cytoplasm In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. ...
of infected cells and are formed early in the infection cycle. The number and the size of viroplasms depend on the virus, the virus isolate, hosts species, and the stage of the infection. For example, viroplasms of mimivirus have a similar size to the nucleus of its host, the
amoeba An amoeba (; less commonly spelled ameba or amœba; plural ''am(o)ebas'' or ''am(o)ebae'' ), often called an amoeboid, is a type of cell or unicellular organism with the ability to alter its shape, primarily by extending and retracting pseudop ...
''
Acanthamoeba ''Acanthamoeba'' is a genus of amoebae that are commonly recovered from soil, fresh water, and other habitats. ''Acanthamoeba'' has two evolutive forms, the metabolically active trophozoite and a dormant, stress-resistant cyst. Trophozoites are ...
polyphaga''. A virus can induce changes in composition and organization of host cell cytoskeletal and membrane compartments, depending on the step of the
viral replication Viral replication is the formation of biological viruses during the infection process in the target host cells. Viruses must first get into the cell before viral replication can occur. Through the generation of abundant copies of its genome an ...
cycle. This process involves a number of complex interactions and signaling events between viral and host cell factors. Viroplasms are formed early during the infection; in many cases, the cellular rearrangements caused during virus infection lead to the construction of sophisticated inclusions —viroplasms— in the cell where the factory will be assembled. The viroplasm is where components such as replicase enzymes, virus genetic material, and host proteins required for replication concentrate, and thereby increase the efficiency of replication. At the same time, large amounts of ribosomes, protein-synthesis components, protein folding chaperones, and
mitochondria A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is used ...
are recruited. Some of the membrane components are used for viral replication while some others will be modified to produce viral envelopes, when the viruses are enveloped. The viral replication,
protein synthesis Protein biosynthesis (or protein synthesis) is a core biological process, occurring inside cells, balancing the loss of cellular proteins (via degradation or export) through the production of new proteins. Proteins perform a number of critical ...
and assembly require a considerable amount of energy, provided by large clusters of mitochondria at the periphery of viroplasms. The virus factory is often enclosed by a membrane derived from the
rough endoplasmic reticulum The endoplasmic reticulum (ER) is, in essence, the transportation system of the eukaryotic cell, and has many other important functions such as protein folding. It is a type of organelle made up of two subunits – rough endoplasmic reticulum ( ...
or by cytoskeletal elements. In animal cells, virus particles are gathered by the
microtubule Microtubules are polymers of tubulin that form part of the cytoskeleton and provide structure and shape to eukaryotic cells. Microtubules can be as long as 50 micrometres, as wide as 23 to 27  nm and have an inner diameter between 1 ...
-dependent aggregation of toxic or misfolded protein near the
microtubule organizing center The microtubule-organizing center (MTOC) is a structure found in eukaryotic cells from which microtubules emerge. MTOCs have two main functions: the organization of eukaryotic flagella and cilia and the organization of the mitotic and meiotic spind ...
(MTOC), so the viroplasms of animal viruses are generally localized near the MTOC. MTOCs are not found in plant cells. Plant viruses induce the rearrangement of membranes structures to form the viroplasm. This is mostly shown for plant
RNA virus An RNA virus is a virusother than a retrovirusthat has ribonucleic acid ( RNA) as its genetic material. The nucleic acid is usually single-stranded RNA (ssRNA) but it may be double-stranded (dsRNA). Notable human diseases caused by RNA virus ...
es.


Functions

Viroplasm is the location within the infected cell where
viral replication Viral replication is the formation of biological viruses during the infection process in the target host cells. Viruses must first get into the cell before viral replication can occur. Through the generation of abundant copies of its genome an ...
and assembly take place. Wrapping the viroplasm with a membrane, concentrates the viral components required for the
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ...
replication and the morphogenesis of new virus particles, so it increases the efficiency of the processes. The recruitment of cellular membranes and cytoskeleton to generate virus replication sites can also benefit viruses in other ways. Disruption of cellular membranes can, for example, slow the transport of immunomodulatory proteins to the surface of infected cells and protect against innate and acquired immune responses, and rearrangements to cytoskeleton can facilitate virus release. The viroplasm could also prevent virus degradation by
protease A protease (also called a peptidase, proteinase, or proteolytic enzyme) is an enzyme that catalyzes (increases reaction rate or "speeds up") proteolysis, breaking down proteins into smaller polypeptides or single amino acids, and spurring the ...
s and
nuclease A nuclease (also archaically known as nucleodepolymerase or polynucleotidase) is an enzyme capable of cleaving the phosphodiester bonds between nucleotides of nucleic acids. Nucleases variously effect single and double stranded breaks in their t ...
s. In the case of the
Cauliflower mosaic virus Cauliflower mosaic virus (CaMV) is a member of the genus ''Caulimovirus'', one of the six genera in the family ''Caulimoviridae'', which are pararetroviruses that infect plants. Pararetroviruses replicate through reverse transcription just like ...
(CaMV), viroplasms improve the virus transmission by the
aphid Aphids are small sap-sucking insects and members of the superfamily Aphidoidea. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white woolly aphids. A t ...
vector Vector most often refers to: *Euclidean vector, a quantity with a magnitude and a direction *Vector (epidemiology), an agent that carries and transmits an infectious pathogen into another living organism Vector may also refer to: Mathematic ...
. Viroplasms also control release of virions when the insect stings an infected plant cell or a cell near the infected cells.


Possible co-evolution with the host

Aggregated structures may protect viral functional complexes from the cellular degradation systems. For example, formation of viral factories of the ASFV viroplasm is very similar to the aggresome formation. An
aggresome In eukaryotic cells, an aggresome refers to an aggregation of misfolded proteins in the cell, formed when the protein degradation system of the cell is overwhelmed. Aggresome formation is a highly regulated process that possibly serves to organize m ...
is a perinuclear site where misfolded
proteins Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
are transported and stored by the cell components for their destruction. It has been proposed that the viroplasm could be the product of a co-evolution between the virus and its host. It is possible that a cellular response originally designed to reduce the toxicity of misfolded proteins is exploited by cytoplasmic viruses to improve their replication, the virus
capsid A capsid is the protein shell of a virus, enclosing its genetic material. It consists of several oligomeric (repeating) structural subunits made of protein called protomers. The observable 3-dimensional morphological subunits, which may or ma ...
synthesis, and assembly. Alternatively, the activation of host defense mechanisms may involve sequestration of virus components in aggregates to prevent their dissemination, followed by their neutralisation. For example, viroplasms of mammalian viruses contain certain elements of the cellular degradation machinery which might enable cellular protective mechanisms against viral components. Given the co-evolution of viruses with their host cells, changes in cell structure induced during infection are likely to involve a combination of the two strategies.


Use in diagnostics

Presence of viroplasms is used to diagnose certain viral infections. Understanding the phenomena of virus aggregation and of the cell response to the presence of virus, and whether viroplasms facilitate or inhibit viral replication, may help to develop new therapeutic approaches against virus infections in animal and plant cells.


See also

*
Viral evolution Viral evolution is a subfield of evolutionary biology and virology that is specifically concerned with the evolution of viruses. Viruses have short generation times, and many—in particular RNA viruses—have relatively high mutation rates (on the ...


References

{{Portal bar, Viruses Virology