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1 The Leslie and Susan Gonda (Goldschmied) Multidisciplinary Brain Research Center, Bar Ilan University, Ramat Gan, Israel; Faculty of Life Sciences, Bar Ilan University, Ramat Gan, Israel
2 Faculty of Life Sciences, Bar Ilan University, Ramat Gan, Israel
3 The Leslie and Susan Gonda (Goldschmied) Multidisciplinary Brain Research Center, Bar Ilan University, Ramat Gan, Israel
* To whom correspondence should be addressed. E-mail: avy{at}mail.biu.ac.il.
Inhibiting NO synthesis during learning that food is inedible in Aplysia blocks subsequent memory formation. To gain insight into the function of NO transmission during learning we tested whether blocking NO synthesis affects aspects of feeding that are expressed in both a non-learning context as well as during learning. Inhibiting NO synthesis with L-NAME and blocking guanylyl cyclase with methylene blue decreased the efficacy of ad libitum feeding. D-NAME had no effect. L-NAME also decreased rejection responses frequency, but did not affect rejection amplitude. The effect of L-NAME was explained by a decreased feedback when efforts to swallow are not successful, leading to a decreased rejection rate, and a decreased ability to re-position and subsequently consume food in ad libitum feeding. Feedback from efforts to swallow are a critical component of learning that food is inedible. Treatment with the NO donor SNAP substituted for food entry into the mouth and efforts to swallow, thereby converting an experience that does not produce memory, lip stimulation alone, into an experience producing memory. Exogenous NO at a concentration causing memory also excited a key neuron responding to NO, the MCC. Block of the cGMP second messenger cascade during training via methylene blue also blocked memory formation after learning. Our data indicate that memory arises from the contingency of three events during learning that food is inedible. One of the events is efforts to swallow, which are signaled by NO via cGMP.
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