Mosaic disease

Tobacco mosaic virus
Electron micrograph of TMV particles stained to enhance visibility at 160,000x magnification
Virus classification
Group:	Group IV ((+)ssRNA)
Order:	Unassigned
Family:	Virgaviridae
Genus:	Tobamovirus
Species:	Tobacco mosaic virus

Tobacco mosaic virus coat protein
A monomeric unit of the tobacco mosaic virus coat protein.
Identifiers
Symbol	CP
Entrez	1494073
UniProt	P03579

Tobacco mosaic virus (TMV) is a positive-sense single stranded RNA virus that infects a wide range of plants, especially tobacco and other members of the family Solanaceae. The infection causes characteristic patterns, such as "mosaic"-like mottling and discoloration on the leaves (hence the name). TMV was the first virus ever to be discovered. Although it was known from the late 19th century that an infectious disease was damaging tobacco crops, it was not until 1930 that the infectious agent was determined to be a virus.

In 1886, Adolf Mayer first described the tobacco mosaic disease that could be transferred between plants, similar to bacterial infections. In 1892, Dmitri Ivanovsky gave the first concrete evidence for the existence of a non-bacterial infectious agent, showing that infected sap remained infectious even after filtering through finest Chamberland filters. Later, in 1903, Ivanovsky published a paper describing abnormal crystal intracellular inclusions in the host cells of the affected tobacco plants and argued the connection between these inclusions and the infectious agent. However, Ivanovsky remained rather convinced, despite repeated failures to produce evidence, that the causal agent was an unculturable bacterium, too small to be retained on the employed Chamberland filters and to be detected in the light microscope. In 1898, Martinus Beijerinck independently replicated Ivanovsky's filtration experiments and then showed that the infectious agent was able to reproduce and multiply in the host cells of the tobacco plant. Beijerinck coined the term of "virus" to indicate that the causal agent of tobacco mosaic disease was of non-bacterial nature. Tobacco mosaic virus was the first virus to be crystallized. It was achieved by Wendell Meredith Stanley in 1935 who also showed that TMV remains active even after crystallization. For his work, he was awarded 1/3 of the Nobel Prize in Chemistry in 1946, even though it was later shown some of his conclusions (in particular, that the crystals were pure protein, and assembled by ) were incorrect. The first electron microscopical images of TMV were made in 1939 by Gustav Kausche, Edgar Pfankuch and Helmut Ruska – the brother of Nobel Prize winner Ernst Ruska. In 1955, Heinz Fraenkel-Conrat and Robley Williams showed that purified TMV RNA and its capsid (coat) protein assemble by themselves to functional viruses, indicating that this is the most stable structure (the one with the lowest free energy). The crystallographer Rosalind Franklin worked for Stanley for about a month at Berkeley, and later designed and built a model of TMV for the 1958 World's Fair at Brussels. In 1958, she speculated that the virus was hollow, not solid, and hypothesized that the RNA of TMV is single-stranded. This conjecture was proven to be correct after her death and is now known to be the + strand. The investigations of tobacco mosaic disease and subsequent discovery of its viral nature were instrumental in the establishment of the general concepts of virology.