Long-term memory and response generalization in mushroom body extrinsic neurons in the honeybee Apis mellifera

Honeybees learn to associate an odor with sucrose reward under conditions that allow the monitoring of neural activity by imaging Ca(2+) transients in morphologically identified neurons. Here we report such recordings from mushroom body extrinsic neurons - which belong to a recurrent tract connecting the output of the mushroom body with its input, potentially providing inhibitory feedback - and other extrinsic neurons. The neurons' responses to the learned odor and two novel control odors were measured 24 h after learning. We found that calcium responses to the learned odor and an odor that was strongly generalized with it were enhanced compared with responses to a weakly generalized control. Thus, the physiological responses measured in these extrinsic neurons accurately reflect what is observed in behavior. We conclude that the recorded recurrent neurons feed information back to the mushroom body about the features of learned odor stimuli. Other extrinsic neurons may signal information about learned odors to different brain regions.