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The key belonged to the Men of Letters and was kept in a box which was decorated with their crest — the Aquarian Star. According to Larry Ganem, inside the box is “the key to every object, scroll, and spell ever collected, for thousands of years, under one roof.” The Men of Letters had stored, in one location, all the supernatural knowledge that they had ever possessed. If the Box were to be hunted down by any “evil being” (demons, monsters, etc…) it would be safe when stored in the area where they had placed all their scrolls and secrets because the area itself is branded with symbols to ward off every kind of evil ever created.
It fell into the possession of the Winchesters after their grandfather, Henry Winchester, died at the hands of a demon, Abaddon, while the demon was out hunting for it to obtain the Bunker’s secret knowledge.
According to British Men of Letters operative Mick Davies, the key actually opens every Men of Letters chapter house in the world, not just the American Bunker.
In 1958, Abaddon, one of the Knights of Hell, kills the members of the group who were gathered in Normal, Illinois. They were gathered for the initiation of Henry Winchester and Josie Sands.
Abaddon killed David Ackers and Ted Bowen, and blinded Larry Ganem. After Henry bursts through to find out what was happening, Larry crawls up to Henry and entrusts him with a box which contains the key to where all the knowledge the Men of Letters has collected is stored. Henry goes through a portal created by his blood spell, and is followed by Abbadon.
Henry meets up with Sam and Dean, and together they manage to track down the sole survivor of the massacre.
Sam visits Larry Ganem to find out more about the box and its contents. After Larry explains everything, Abaddon, who has possessed Larry’s wife and was listening to the entire conversation, kills Larry and captures Sam. Abaddon then calls Dean to demand a trade: Sam for Henry & the key. During the trade, Henry shoots Abaddon in the head with a bullet that had a devil’s trap carved into it. This immobilizes Abaddon where she stands, and she screams “Why am I stuck?!” in frustration. Dean then decapitates her and says that he will cut her up into strips and bury her. After Henry’s death, Sam and Dean decide to find out if the Bunker is still intact.
Sam and Dean follow the coordinates provided by Larry, and find an underground Bunker, which they are able to enter using the key.
Dean hands Kevin the key to the Men of Letters Bunker where he can be safely protected.
In 1935, two Men of Letters, James Haggerty and Peter Jenkins use the key to enter the recently finished Bunker
Sam and Dean flash the box holding the key when they are trying to get Cuthbert Sinclair’s attention outside of his magically hidden mansion.
Lady Antonia Bevell uses a copy of the key to gain entry into the Men of Letters Bunker to lay a trap for Sam, Dean and Castiel.
Mick Davies told Sam and Dean that the key to the bunker can open every Men of Letters chapter house scattered across the world.
After arriving at the Men of Letters’ Rhode Island Chapter, Sam and Dean find a man-hole cover with the Aquarian Star in the center and a key hole. Sam uses the key to open the hatch and gain access to the hallways beneath the building.