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Very interesting game! I included it in my compilation video series of all the games entered in the GameMaker Studio 2 Beta Jam #2, if you'd like to take a look :)
Some cool ideas here, and a coherent effective art style + atmosphere. Learning how to set up your first runes and towers and work out what they do or how effective they are takes a long time though, and this sort of "lots to learn" is a problem of many tower defense games already!
Perhaps some single rune attack towers that start the player off, and reserve the more complex arrangements for more and more powerful structures would give the player more time to learn the systems.
I did some attempt, placed down the runes according to the information in top left; but it didn't seem to do that much. So a bit confusing. But the game definatly looks very nice and interesting; like a sort of different take on a tower defense game, which is cool! Good luck!