I think that this observation was posted before, but cannot find it. So here it is again.
The big right-hand flower on page You are not allowed to view links.
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Login to view. extends for a few mm into the other half of the bifolio, f40r; proving that the
drawings were done while the bifolios were still unbound.
That flower has been proposed as evidence that the
color paints too were applied before the bifolios were bound, since the bits of the flower on You are not allowed to view links.
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However, looking closely, on You are not allowed to view links.
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So it seems that the book was already bound when that blue paint was applied. The Painter must have noticed that the flower of You are not allowed to view links.
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The blue paint is distinguished from other paints for frequently producing dark gray "offset" (contact) prints on facing pages. The prints presumably were due to some component of the blue paint, not the blue pigment itself, that did not dry properly. Here it is no exception: we see offsets of You are not allowed to view links.
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Those offset prints were obviously produced when the bifolios were already nested in the current order. Moreover, their vertical positions precisely match those of the original painted aeras; so it is highly likely that the bifolios were already bound, not simply nested but loose, when the offsetting happened.
It is possible that the paint was applied when the bifolios were loose, but the offsetting started only centuries after that, after the book was bound. Maybe because the book was kept for a while in a damp environment and the humidity softened that dark gray component of that particular paint.
But it seems more likely that the offset prints formed right after the paint was applied, because the paint included an improper ingredient that did not dry properly -- like tar, perhaps added to get a darker shade of blue. Indeed, the mottled appearance of the dark blue paint could be explained as due to some oily component that did not mix well with the water-based binder. Or maybe all the dark blue areas were painted in a single pass, and the Painter did not wait for the paint to dry before putting the two sheets in contact and moving on to the next drawing. In that case, the offset prints on You are not allowed to view links.
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Login to view. and f39v would be evidence that the paint was applied after the binding -- that is, long after the outlines were drawn.
All the best, --stolfi