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In Desktop Defender, you can either upgrade a tower by spending money in game or by spending coins out of game (in the metagame). When do the out of game upgrades take effect? Do they affect level 1 towers, or do you have to upgrade those towers in game before they do anything?

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You have to upgrade those towers in game; what the out-of-game upgrades do is basically unlock in-game tower ranks, but you must still upgrade in-game to the new rank.

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