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. 2011 Mar 22;278(1707):898-905.
doi: 10.1098/rspb.2010.1891. Epub 2010 Nov 10.

Conceptualization of above and below relationships by an insect

Aurore Avarguès-Weber  1 , Adrian G Dyer, Martin Giurfa
Affiliations

Conceptualization of above and below relationships by an insect

Aurore Avarguès-Weber et al. Proc Biol Sci. .

Abstract

Relational rules such as 'same' or 'different' are mastered by humans and non-human primates and are considered as abstract conceptual thinking as they require relational learning beyond perceptual generalization. Here, we investigated whether an insect, the honeybee (Apis mellifera), can form a conceptual representation of an above/below spatial relationship. In experiment 1, bees were trained with differential conditioning to choose a variable target located above or below a black bar that acted as constant referent throughout the experiment. In experiment 2, two visual stimuli were aligned vertically, one being the referent, which was kept constant throughout the experiment, and the other the target, which was variable. In both experiments, the distance between the target and the referent, and their location within the visual field was systematically varied. In both cases, bees succeeded in transferring the learned concept to novel stimuli, preserving the trained spatial relation, thus showing an ability to manipulate this relational concept independently of the physical nature of the stimuli. Absolute location of the referent into the visual field was not a low-level cue used by the bees to solve the task. The honeybee is thus capable of conceptual learning despite having a miniature brain, showing that such elaborated learning form is not a prerogative of vertebrates.

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Figures

Figure 1.
Figure 1.
Stimuli used in the experiments. (a) Experiment 1: during training, the targets were presented above and below the referent bar. (b) Experiment 2: only achromatic patterns were used. During training, the targets were presented above and below the referent pattern that was a cross or a disc depending on the group of bees trained.
Figure 2.
Figure 2.
Experiment 1. (a) Example of the conditioning and testing procedure. Half of the bees were rewarded on the ‘target above bar’ relation whereas the other half was rewarded on the ‘target below bar’ relation. The transfer test was not rewarded. (b) Acquisition curve during training (percentage of correct choices as a function of blocks of five trials) and performance (cumulative choices during 45 s test) in the non-rewarded transfer test (white bar). Data shown are means and s.e.m. (n = 8). Bees succeeded in learning the rule based on the above versus below relationship and in transferring the concept to novel stimuli (*p < 0.05; ***p < 0.001).
Figure 3.
Figure 3.
Experiment 2. (a) Example of the conditioning and testing procedure. Half of the bees were rewarded on the ‘target above referent’ relation whereas the other half was rewarded on the ‘target below referent’ relation. The referent pattern was either the disc or the cross depending on the group of bees trained. The transfer test was not rewarded. (b) Phase 1 (pre-training): acquisition curve during pre-training (percentage of correct choices as a function of blocks of five trials) and performance (cumulative choices during 45 s test) in the non-rewarded discrimination test (white bar). Data shown are means and s.e.m. (n = 20). Bees learned to choose the referent pattern and to discriminate it from other patterns used as targets in the subsequent training phase (**p < 0.01; ***p < 0.001). (c) Phase 2 (training): acquisition curve during training (percentage of correct choices as a function of blocks of five trials) and performance (cumulative choices during 45 s test) in the non-rewarded tests (white bars). Data shown are means and s.e.m. (n = 20 for acquisition curve and transfer test, and n = 8 for controls 1 and 2). The inset shows acquisition performance during the first five trials that integrate the first training block. Bees learned the concept of above/below and transferred it to novel stimuli. Controls 1 and 2 show that the spatial location of the referent on the background was not used as a discrimination cue to resolve the task (*p < 0.05; **p < 0.01; ***p < 0.001).
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