Tobacco mosaic virus (TMV) is an ​RNA virus​ that infects ​plants​, especially

tobacco​ and other members of the family ​Solanaceae​. The ​infection​ causes
characteristic patterns (​mottling​ and discoloration) on the ​leaves​ (thus the
name). TMV was the first ​virus​ to be discovered. Although it was known from
the late 19​th​ ​century​ that an ​infectious disease​ was damaging tobacco crops, it
was not until ​1930​ that the infectious agent was determined to be a virus.
History

In ​1883​, ​Adolf Mayer​ first described the disease that could be transferred between plants, similar
to ​bacterial​ infections​[1]​. ​Dimitri Ivanovski​ gave the first concrete evidence for the existence of a
non-bacterial infectious agent, showing that infected sap remained infectious even after filtering
through ​Chamberland filter​ ​candles​, in ​1892​. However, he remained convinced despite repeated
failures to produce evidence, that bacteria were the infectious agents. In 1898, ​Martinus
Beijerinck​ showed that a filtered, ​bacteria​-free culture medium still contained the infectious
agent​[1]​. ​Wendell Meredith Stanley​ crystallized the virus in ​1935​ and showed that it remains
active even after crystallization​[1]​. For his work, he was awarded 1/4 of the ​Nobel Prize in
Chemistry​ in ​1946​[2]​, even though it was later shown some of his conclusions (in particular, that
the crystals were pure protein, and assembled by ​autocatalysis​) were incorrect.​[3]​ 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​.[4]​ ​ 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), and likely the natural assembly mechanism
within the ​host cell​.

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.

[​edit​] Structure
Structure
Schematic model of TMV: 1. nucleic acid (​RNA​), 2. capsomer (protomer), 3. ​capsid​.

Electron micrograph of TMV particles stained to enhance visibility at 160,000x magnification.

Tobacco mosaic virus has a rod-like appearance. Its capsid is made from 2130 ​molecules​ of coat
protein (see image above) and one molecule of genomic RNA 6390 bases long. The coat protein
self-assembles into the rod like helical structure (16.3 proteins per helix turn) around the RNA
which forms a hairpin loop structure (see the ​Electron Micrograph​ below). The protein monomer
consists of 158 ​amino acids​ which are assembled into four main alpha-helices, which are joined
by a prominent loop proximal to the axis of the virion. Virions are ~300 nm in length and ~18
nm in diameter.​[5]​ Negatively stained electron microphotographs show a distinct inner channel of
~4 nm. The RNA is located at a radius of ~6 nm and is protected from the action of cellular
enzymes by the coat protein. There are three RNA nucleotides per protein monomer.​[6]​ TMV is a
thermostable virus. On a dried leaf, it can withstand upto 120 degrees for 30 minutes

[​edit​] Replication

When it enters the host, it enters as single stranded RNA and has a temporary double stranded
intermediate. Its RNA directs the synthesis of viral proteins by the host machinery. After the
RNA and the protein is produced, they undergo self assembly. After the assembly of capsomeres
and RNA, they are released outside the cell after the ​lysis​ and death of the host. These newly
synthesized viruses continue on to infect other cells.

[​edit​] Infection

When TMV infects a tobacco plant, the virus enters mechanically (For example through a
ruptured plant cell wall) and replicates. After its multiplication, it enters the neighboring cells
through ​plasmodesmata​. For its smooth entry, TMV produces a 30,000 ​Dalton​ protein called P30
which tends to enlarge the plasmodesmata. TMV most likely moves from cell-to-cell as a
complex of the RNA, P30, and replicase proteins. The first symptom of this virus disease is a
light green coloration between the veins of young ​leaves​. This is followed quickly by the
development of a “mosaic” or mottled pattern of light and dark green areas in the leaves. These
symptoms develop quickly and are more pronounced on younger leaves. Mosaic does not result
in plant death, but if infection occurs early in the season, plants are stunted. Lower leaves are
subjected to “mosaic burn” especially during periods of hot and dry weather. In these cases, large
dead areas develop in the leaves. This constitutes one of the most destructive phases of tobacco
mosaic virus infection. Infected leaves may be crinkled, puckered, or enlongated.

[​edit​] Scientific and Environmental Impact

In plants, tobacco mosaic virus leads to severe crop losses​[7]​. It is known to infect members of
nine plant families, and at least 125 individual species, including tobacco, tomato, pepper,
cucumbers, and a number of ornamental flowers.​[8]​ There are many different strains.

The large amount of literature about TMV and its choice for many pioneering investigations in
structural molecular biology​, ​X-ray diffraction​, virus assembly and disassembly, and so on, are
fundamentally due to the large quantities that can be obtained, plus the fact that it does not infect
animals. After growing a few infected tobacco plants in a ​greenhouse​ and a few simple
laboratory procedures, a scientist can easily produce several grams of virus. As a result of this,
TMV can be treated almost as an organic chemical, rather than an infective agent.