reference of which

Are both of the highlighted "which"'s referring to "images" or "circuits of neurons"?

Context:

The neuroscientist Antonio Damasio explains that one of the brain’s main tasks is continually to monitor and regulate our bodily functions, and it does this through establishing self-reflexive maps – images linking circuits of neurons which are established in the brain to correspond to different bodily activities, between which there is two-way feedback. This is essential to our well-being and obviously applies to routine physical functions such as the monitoring of body temperature, or the release of hormones to increase appetite stimulated by a fragrant waft of cooking, for example.