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For several decades, the Caspar-Klug theory faithfully described all the known viral structures. But with the development of higher-resolution imaging techniques like cryo-electron microscopy, exceptions soon appeared to the strict constraints on the virus capsid composition (60T subunits) predicted by the theory. The first detected outliers were the cancer-causing polyoma and papilloma viruses, which have capsids made up of 72 pentamer facets.
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