Proofs are a different animal entirely. All of the 1st issue revenue proofs are much brighter in color than the actual stamps (actually, the same with front-of-book proofs). The highest quality ink, paper, and care and production, whereas the actual stamps... well, not so much.
There may be other reasons. Admittedly, proofs are not my area.
Over time, the lighter colored stamps tend to get darker and the dark colored stamps tend to get lighter. This was a deliberate attempt to deal with the cleaning and reusing of the stamps, which was a serious problem and eventually led to the second and third issues being printed.
Proofs were printed with new plates on India paper, which is a very hard, thin, white paper which takes great impressions, or on thin card, again very white, which also takes great impressions.
I collect plate varieties on proofs and can testify that proofs were sometimes taken in both the early and late states. The Smithsonian has proof sheets of R34 10 cent Contract in both early and late states. I have a late state proof sheet of R115. I have seen the " double transfer throughout" of R 24 P4, sorry I didn't buy it. Proofs also had the advantage of not being glued to a piece of paper and stuffed into a shoebox. I suspect proofs were taken on a regular basis throughout the printing run, although Deats claimed to have lost a number of sheets to mice and squirrels in his barn.
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