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the easy answer is "everything"
even if you don't have any gardens/babywithbeard/snipergun/etc, you probably have a ton of enemies that have multiple outline shaders on them (icecube/curse etc)
could be anything. godot actually has good builtin profiling tools for this stuff, but if i had to guess there isn't a lot of incentive to optimize the game for long endless runs ... it would be ultimately pointless and a lot like playing endless but for the developers: there's always a limit to reach!
does the whole game run single threaded for everyone, or is that just under proton/wine on linux?
Everything makes a sound, this makes lag.
Everything is animated, this makes lag.
Projectiles everywhere, they all have to be tracked, there's 10+ events per projectile.
"AI" has to be calculated every microsecond.
Character input has to be calculated every microsecond.
All the % chance items and stats have to be calculated for every event.
Special effects everywhere.
Creation of items on the stage creates lag.
Using arrays instead of collections creates lag now...
Game engines are not as optimized as they need to be, then a developer doing stuff with the engine just reduces the performance.
All of the options in the settings do absolutely nothing but reduce visual clutter, they don't remove anything. All the things that reduce opacity don't stop those special effects from happening when set to zero, they just don't get pushed to the monitor, they still have to be rendered. They should probably be turned OFF and removed from the render pipeline instead.