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Cowpea Chlorotic Mottle Virus, known by the abbreviation CCMV, is a virus that specifically infects the cowpea plant, or black-eyed pea. The leaves of infected plants develop yellow spots, hence the name "chlorotic". Similar to its "brother" virus, Cowpea mosaic virus (CPMV), CCMV is produced in high yield in plants. In the natural host, viral particles can be produced at 1–2 mg per gram of infected leaf tissue. Like the majority of viruses, CCMV consists of a protein shell, that surrounds a central core of RNA. Its protein shell, or capsid, is composed of icosahedral arrangements of identical protein subunits. The symmetry at which this is done is exactly like the symmetry of a football: hexagons and pentagons are interchanged to give the final sphere. The virus is about 30 nm in size, and can be observed with transmission electron microscopy using negative staining.

History

Bancroft et al. in 1967 described the first experiments to isolate and characterise the virus. Since that time, due to the relative ease with which it is grown and isolated, many researchers have focused their attention on the virus. The interest of the scientific community for this virus is also due to a conspicuous property: it is possible to disassemble the virus and remove the genetic material, the RNA. Then, under slightly acidic pH and with relatively high amounts of salts, it is possible to stimulate the self-assembly of the protein subunits, into a shell of identical size to the virus. This yields an empty capsid which has a number of interesting properties. Several successful attempts are reported to incorporate other materials, such as inorganic crystals, inside the capsid.

The promise of CCMV in nanotechnology

CCMV has a number of physical properties that can be exploited for nanoscale fabrications. As previously stated, viruses consist of a protein coat, called a capsid, and nucleic acid (either RNA or DNA). The capsid's main function is to protect the nucleic acid. Mutagenesis of viral capsids is a well-established technique that permits the alteration of the capsid surface. Such alterations can include changing the charge or attaching specific ligands to the capsid surface via their chemical interactions with surface molecules or amino acids. Viral capsids have been shown to allow the attachment of small peptides (<30 amino acids) onto their surface. Furthermore, heterologous expression of CCMV in yeast allows large-scale production of wild-type or genetically modified CCMV capsids. These capsids can either contain or be empty of nucleic acid. Without nucleic acid, these cage-like structures provide cavities of well-defined size, shape, and charge, and can be used for particle-specific entrapment of organic and inorganic materials.

Some of the physical properties of CCMV are: toleration of high temperatures, a variety of pH's, and stability in organic solvents, such as DMSO. These conditions allow for a wide range of chemistry to be tolerated in CCMV modification. One of the most profound characteristics of CCMV is that the viral capsid can undergo reversible pH and metal-ion structural transition, without any loss of viral function or structural damage. This transition, referred to as "swelling," forms 60 separate 2 nm sized openings in the protein shell. Opening of the cavity occurs under slightly acidic conditions at a pH of about 6.5. The capsule can be closed by reversing the pH to about 5.0. The swollen capsid permits ions to diffuse freely into and out of the cavity. Entrapment of organic or inorganic materials is only restricted by the interactions of these molecules with molecules of the viral interior. The interior and exterior properties of the viral capsid are dissimilar enough to control the localization of molecules within the protein cage. The cationic interior would naturally encapsulate anionic species, but the charge of the cavity can be altered by site-directed mutagenesis.

Additionally, accessible amines and carboxylates representing potential sites for external modification have been identified on the exterior surface of the viral capsid. These groups can be modified to facilitate specific biological targeting or surface interactions through the linking of various ligands. Fluorescent dyes have also been shown to be selectively attached to the capsid surface.

As the virus' natural ability to infect host cells can be exploited, the use of CCMV is of novel interest to researchers. CCMV has feasible applications in biosensors, nanoelectric devices, and drug targeting and delivery. One recent example shows how DNA origami nanostructures can be coated by CCMV capsid proteins and subsequently, efficiently delivered into human cells.^[1]

Lastly, though CPMV exhibits similar physical properties to CCMV, CCMV has been shown to be more easily modified, less susceptible to degradation, and is not shown to be specific for particular mammalian cells in vivo.

References

^ J. Mikkilä, A.-P. Eskelinen, E. H. Niemelä, V. Linko, M. J. Frilander, P. Törmä and M. A. Kostiainen.Nano Lett.,14(4),2196 (2014)

CCMV Overview
Bancroft JB, Hiebert E. Formation of an infectious nucleoprotein from protein and nucleic acid isolated from a small spherical virus. Virology, 1967
Mikkilä, J. et al. Virus-Encapsulated DNA Origami Nanostructures for Cellular Delivery, "Nano Lett." 2014 [1]
Douglas, T. & Young, M. Host–guest encapsulation of materials by assembled virus cages, Nature, 1998 [2]
Destito, G. et al. Folic-acid mediated targeting of Cowpea mosaic virus particles to tumor cells. Cell Press, 2007
Steinmetz, N. & Evans, D.J. Utilization of plant viruses in bionanotechnology, Organic and Biomolecular Chemistry, 2007

[3]

External links

ICTVdB - The Universal Virus Database: Cowpea chlorotic mottle virus
Family Groups - The Baltimore Method

English Journal

A novel set of polyvalent primers that detect members of the genera Bromovirus and Cucumovirus.

Seo JK1, Lee YJ1, Kim MK1, Lee SH2, Kim KH3, Choi HS4.
Journal of virological methods.J Virol Methods.2014 Jul;203:112-5. doi: 10.1016/j.jviromet.2014.03.026. Epub 2014 Apr 6.
Rapid detection and diagnosis of plant virus infection is one of the most important steps in preventing damages caused by viral diseases. Bromoviruses and cucumoviruses belong to the family Bromoviridae, which is one of the most important families of plant viruses, and infect a broad range of host p
PMID 24717165

Chemotherapy pro-drug activation by biocatalytic virus-like nanoparticles containing cytochrome P450.

Sánchez-Sánchez L1, Cadena-Nava RD2, Palomares LA1, Ruiz-Garcia J3, Koay MS4, Cornelissen JJ4, Vazquez-Duhalt R5.
Enzyme and microbial technology.Enzyme Microb Technol.2014 Jun 10;60:24-31. doi: 10.1016/j.enzmictec.2014.04.003. Epub 2014 Apr 13.
This work shows, for the first time, the encapsulation of a highly relevant protein in the biomedical field into virus-like particles (VLPs). A bacterial CYP variant was effectively encapsulated in VLPs constituted of coat protein from cowpea chlorotic mottle virus (CCMV). The catalytic VLPs are abl
PMID 24835096

Versatile post-functionalization of the external shell of cowpea chlorotic mottle virus by using click chemistry.

Hommersom CA1, Matt B, van der Ham A, Cornelissen JJ, Katsonis N.
Organic & biomolecular chemistry.Org Biomol Chem.2014 May 28;12(24):4065-9. doi: 10.1039/c4ob00505h.
We present the modification of the outer protein shell of cowpea chlorotic mottle virus (CCMV) with linear and strained alkyne groups. These functionalized protein capsids constitute valuable platforms for post-functionalization via click chemistry. After modification, the integrity of the capsid an
PMID 24817149

Japanese Journal

(64)植物トランジェント発現系を利用したブロモウイルス複製酵素の解析

藤崎恒喜,海道真典,三瀬和之,奥野哲郎
日本植物病理學會報 69(1), 68, 2003-02-25
… chlorotic mottle virus(CCMV)は互いに近縁なブロモウイルスである.ブロモウイルスのRNA1と2にはそれぞれウイルスRNA複製酵素の構成成分である1a,2aタンパク質がコードされている.Nicotiana benthamianaのプロトプラストにおいて,BMVやCCMVではRNA2をSBLVのRNA2と交換してもウイルスRNAの蓄積が認められるが,RNA1の交換では認められない.この原因のひとつとしてRNA1またはRNA2の非効率的な増幅による1aまたは2aタンパ …
NAID 110002716780

(334)BMV 3a移行タンパク質遺伝子を持つキメラCCMV適応変異体の解析(平成9年度日本植物病理学会大会講演要旨)

藤田泰成,三瀬和之,古澤巌
日本植物病理學會報 63(3), 274, 1997-06-25
NAID 110007723441

(257) Bromovirusの宿主特異性 (1) : BMVの3a遺伝子をもつハイブリッドCCMV由来のpseudorevertantsのcDNAクローニング (日本植物病理学会大会)

三瀬和之,AHLQUIST P.
日本植物病理學會報 60(3), 385, 1994-06-25
NAID 110002737178

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リンク元	「ササゲクロロティックモトルウイルス」「cowpea chlorotic mottle virus」
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Cowpea chlorotic mottle virus (CCMV)
Virus classification
Group:	Group IV ((+)ssRNA)
Order:	Unassigned
Family:	Bromoviridae
Genus:	Bromovirus
Species:	Cowpea chlorotic mottle virus
Synonyms
	Bean yellow stipple virus; Virus del moteado amarillo;