They are conjugational endings from (sh)i-Adjectives:
We can derive verbs from adjectives by adding :
> -> ->
And conjugate them further:
*
* ()
* (negation)
* ...
This conjugational suffix got reanalyzed as a word on its own, yielding , which was shortened to , , and .
Even by itself was used in the meaning of by the same process.
> _ __ _=