Speed and skill. Each pencil drawing by an animator was traced in ink on to a transparent sheet. Once the ink dried it was sent to painting where the paint was applied to the BACK of the transparent film. Each section of color had to be completed quickly while the paint was still very wet so it would show no brushstrokes or the cel was ruined, but going outside the lines would also ruin a cel, as even though it would not obscure the line, it would still show the color in the next section over.
In the golden age of Disney the ink and paint department was a fleet women who’s only job was to trace or paint. The inking girls were considered to be a higher caliber than the painting girls, as their work required a more steady hand, but the painting girls were amazing in their own right. One of the greatest accomplishments of that department was maintaining consistency in color and positioning on Snow White’s blushed cheeks, which were done with *actual blush.* The effect was so time consuming it was never used again. They stuck to solid colors from then on out.
Edit: I have been corrected below. The use of actual blush is an urban legend. The effect was achieved no less amazingly with a dye applied to each cel by a very talented woman from inking named Helen Ogger. See the post correcting me below for more detail.
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