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11Also at Cambridge, Sydney Brenner and virologist Robert Horne developed a new method of “negative staining” that allowed for more fine-grained electron microscopic imaging of viral capsids, indispensable for later theory development on their structure. In negative staining, it is the imaging background that is stained, leaving the specimens untouched and visible by contrast (while in positive staining, the specimen itself is stained). In electron microscopy, negative staining involves applying a heavy metal salt like uranium acetate or phosphotungstic acid on the imaging grid, so that it strongly scatters electrons and appears dark. The specimen remains relatively electron-transparent and appears bright on the EM images. This method is especially well-suited for naturally low-density specimens like viral capsids.
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