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On-ball memory assay. Bottom, timeline of associative learning and memory assay for flies on a trackball. Each fly had three bouts of odour testing before conditioning (Pre), characterizing its initial responses to the CS + and CS - odours. Next, there were six training bouts, each with a paired presentation of the CS + and the unconditioned stimulus (US, electric shock pulses) plus an unpaired CS - presentation. Finally, each fly had three testing bouts at both 5 min and 1 h after conditioning. Red line indicates the end of training. Top, timelines for individual training and testing bouts. A pair of innately attractive odours (ACV and 1% ethyl acetate) were counterbalanced as CS + and CS - across flies for b - f .

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