To what do the two relative pronouns "which" refer?

In the sentence:

The sense that life as a whole is absurd arises when we perceive, perhaps dimly, an inflated pretension or aspiration which is inseparable from the continuation of human life and which makes its absurdity inescapable, short of escape from life itself.

To what do the two relative pronouns "which" refer here? That is, what is inseparable from life and what makes its absurdity inescapable? The "pretension or apiration" or the "perceiving"? How can one tell here? Would one utilize a comma while the other wouldn’t? Is there a difference between AmE and BrE?

Thanks