Significance
In 1898, Martinus W. Beijerinck, of the Netherlands, put forth his concepts that TMV was small and infectious. Furthermore, he showed that TMV could not be cultured, except in living, growing plants. This report, suggesting that 'microbes' need not be cellular, was to forever change the definition of pathogens. In 1946, Wendall Stanley was awarded the Nobel Prize for his isolation of TMV crystals, which he incorrectly suggested were composed entirely of protein. Research by F.C. Bawden and N. Pirie, in England, during the same period correctly demonstrated that TMV was actually a ribonucleoprotein, composed of RNA and a coat protein. By the mid-1950s, scientists in Germany and the United States proved that the RNA alone was infectious. This discovery ushered in the modern era of molecular virology. TMV is known for several 'firsts' in virology, including the first virus to be shown to consist of RNA and protein, the first virus characterized by X-ray crystallography to show a helical structure (Figure 7), and the first virus used for electron microscopy (Figure 6), solution electrophoresis and analytical ultracentrifugation. TMV also was the first RNA virus genome to be completely sequenced, the source of the first virus gene used to demonstrate the concept of coat protein mediated protection (Figure 11), and the first virus for which a plant virus resistance gene (the N gene) was characterized. Today, TMV is still at the forefront of research leading to new concepts in transgenic technology for virus resistance and developing the virus to act as a 'work horse' to express foreign genes in plants for production of pharmaceuticals and vaccines.