The rune can be upgraded inside the slot though. Note however, that once a rune is placed a special item is required to remove it again. A rune of any level can be placed into an empty slot, there is no need to start with a level 1 rune. Just drag and drop a rune onto the correct slot. Once a slot is unlocked a matching rune can be placed into it. Unlocking a slot on one crusader will not influence the cost to unlock slots on a different crusader. Those unlocking costs are crusader specific. Thus to unlock all 4 currently available slots on a single crusader costs 6 Legendary Catalysts. The cost to unlock the fifth slot is as of yet unknown. The second unlock on a crusader costs 1 Legendary Catalyst, the third costs 2 Legendary Catalysts while the forth unlock costs 3. The first unlock on every crusader is free. Slots can be unlocked in any order on any crusader. Initially all slots on all crusaders are locked and first have to be unlocked to use them. At the bottom of the screen all currently available runes are displayed. The white markers in the little diagrams next to the slot effects are indicating the position of the slot. Those slot effects also are shown when hovering the mouse over a specific slot. Next to it is a list of the Slot Effects for the current crusader. On the right side of the screen the selected crusader with its slots is shown. If you are missing any crusaders in the list this filter likely is the reason. It can be filtered by 'Crusaders in Formation' (the default,) 'Crusaders with Runes' and 'All Crusaders'. On the left side of the screen is a list of crusaders to select from. You can access the rune screen on the same tab that also shows you your trinkets, right next to the button to open the talents, below the button to craft items. So a slot on one crusader may boost DPS while the same slot on another crusader boosts gold find. Every slot comes with an effect, and varies from crusader to crusader. Each crusader has up to 5 different rune slots, though as of this time, all crusaders are limited to 4 slots. Runes ultimately are meant to be equipped on crusaders.
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