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find that it and TMV contain phosphorous conclude that it is not a protein, but is a nucleoprotein TMV TBSV Markham and Smith - 1949 Two classes of particles in purified Turnip yellow mosaic virus preparations: light ones containing only protein, which were not infectious heavy ones containing protein+nucleic acid, which were infectious empty full Myron Brakke- 1951 Development of density gradient centrifugation Isopycnic: particles reach position of equal density in gradient Rate-zonal:Particles sediment differentially through medium as a function of size, shape, and density Equilibrium zonal: Combination of the above Fraenkel-Conrat1955-1956 Virus Aa: RNA A capsid a protein Complete, infectious TMV particles can be reconstituted in vitro from the RNA and protein components RNA alone is infectious RNA can be transcapsidated in protein from closely related virus