Nonelemental visual learning in honeybees

Free-flying honeybees, Apis mellifera, learn visual stimuli in the appetitive context of food search. Visual compound stimuli are relevant in nautre as bees learn flower images that consist of many visual elements. We studied whether elemental associations between each visual element and the reinforcement (elemental approach) are enough to explain the solving of visual discrimination problems that raise ambiguity at the elemental level. We asked whether bees could solve three different visual discriminations: (1) positive patterning (A-, B-, AB+); (2) negative patterning (A+, B+, AB-); and (3) biconditional discrimination (AB+, CD+, AC-, BD-). In experiments 1 and 2 bees had to discriminate a yellow-violet chequerboard from the yellow or the violet squares alone. In experiment 3, four different gratings combining one colour (yellow or violet) with one orientation (vertical or horizontal) had to be discriminated. In all three problems binary compounds were trained in such a way that each element appeared equally often as rewarded and nonrewarded. Bees could solve the three discrimination problems. They always chose the reinforced stimulus despite ambiguity at the level of the elements. For solving positive patterning, elemental processing could be used. For negative patterning and biconditional discrimination, nonelemental processing strategies (unique-cue or configural approach) are necessary to account for these results. Although we cannot decide between a configural and a unique-cue interpretation, we can clearly reject purely elemental processing in these cases. (C) 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.