Effects of fornix transection upon associative memory in monkeys: Role of the hippocampus in learned action

Thirty-three monkeys took part in seven experiments designed to elucidate further the effect of fornix transection on learning and memory. In the first experiment the monkeys had to remember whether stimulus objects had previously been paired with reward or no reward, and they had to use this memory to guide choice between stimulus objects at retention tests according to an arbitrary rule which they had learned: to choose objects previously paired with no reward in preference to objects previously paired with reward. Fornix transection produced a severe and permanent impairment in this task. In the second experiment the monkeys also had to remember object-reward associations but the performance rule was more natural: to choose objects previously paired with reward. Here fornix transection had no effect. The third experiment required the monkeys to remember, given a stimulus object, which of two events of equal valence had previously been the outcome of displacing that object. The two events were either a peanut and a sultana or a black penny and a white penny of equal secondary reinforcing value. Performance was unimpaired by fornix transection. The fourth experiment also demonstrated, in a different paradigm, unimpaired recall of sensory events. The fifth experiment demonstrated an impairment following fornix transection in acquisition of simultaneous spatial-visual conditional discriminations; the sixth demonstrated normal learning by fornix-transected monkeys of a successive spatial-visual conditional discrimination and the seventh demonstrated unimpaired acquisition of a simultaneous auditory-visual conditional discrimination. These results, when considered in detail and together, are incompatible with existing hypotheses of hippocampal function. A new hypothesis is discussed.