Bees travel novel homeward routes by integrating separately acquired vector memories

The question of whether bees can take novel short cuts between familiar sites has been central to the discussion about the existence of cognitive maps in these insects. The failure of bees to show this capacity in the majority of previous studies may be a result of the training procedure, because extensive training to one feeding site may have eliminated or weakened memories for other sites that were previously trained. Here we present a novel approach to this problem, by rewarding honey bees, Apis mellifera carnica, at two feeding sites, one (Sm, 630m, southeast from the hive) at which they could feed in the morning, and the other (Sa, 790m northeast) at which they could feed in the afternoon. We then displaced bees to Sa in the morning and to Sm in the afternoon either from the other feeding site or from the hive. Bees were also displaced to two novel sites, one at a completely unfamiliar location (S4) and another that was located halfway between the two feeding sites (S3). Bees displaced from either of the feeding sites never took novel short cuts; instead, they used the homeward directions that would have been correct had they not been displaced. Bees caught at the hive entrance, however, chose the correct homeward direction not only when displaced to both feeding sites, but also when displaced to S3, although not from S4. Control bees that had been trained to only one of the feeding sites were not able to travel directly home from S3 excluding the possibility that bees used landmarks close to the hive. This is the first evidence that bees take a novel short cut by activating two vector memories simultaneously. The potential mechanisms of integrating the two memories are discussed. Since bees took novel short cuts in only one direction (to the hive) and only when displaced from the hive (not the feeders), we conclude that inference of a cognitive map in bees would be premature.