A creative principle or essence of Ilúvatar, used by him to bring the Ainur into being, and also to bring life into the World of Arda. Tolkien's own definition is found in volume 11 of The History of Middle-earth (Morgoth's Ring): 'This appears to mean the Creative activity of Eru (in some sense distinct from or within Him) by which things could be given real and independent (though derivative and created) existence.'
At first the Flame was with Ilúvatar in the Void, but after the creation of the Ainur, and the Vision of Arda, it was sent to dwell in the heart of the World. By this means, Ilúvatar was able to grant creative power within the World without entering into it Himself. This apparently explains how the Ainur were able to descend into Arda from their own mysterious plane of existence, and how the various rational incarnate races of the World were given independent life.
Melkor was always jealous of the power of the Flame Imperishable, and desired it for himself, but it remained beyond his reach. For this reason, he was unable to create beings of his own, and so his armies were therefore filled with beings he had twisted or corrupted to his own ends.
Actually, it is not entirely clear what happened to the Flame Imperishable at the beginning of the universe. The most direct reference is from Ainulindalë, where Ilúvatar tells the Ainur, 'And I will send forth into the Void the Flame Imperishable, and it shall be at the heart of the World, and the World shall Be; and those of you that will may go down into it.' From the phrasing here it is unclear whether 'the World' means 'the entire universe' or 'the Earth', and other similar passages seem to show the same ambiguity.
Perhaps the most plausible explanation is that this is a creation story in the mythical mould, where the Earth occupies an overwhelmingly central role in the cosmos, so that the two meanings of 'World' are fully intertwined with one another. We see this idea illustrated elsewhere: for example, Varda 'paints' stars onto the sky with the dews of Telperion - a notion that only makes sense in a fully geocentric universe.
If we follow this line of thought, the 'mythical' universe was lost, and the physical came into being, at the time of the Downfall of Númenor, when the West-that-was was taken away from the World, and the World itself was made 'bent' (that is, round). What this profound separation would have meant for the Flame Imperishable, we are not told.
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