Through the looking glass: how do marked dolphins use mirrors and what does it mean?

Mirror-guided self-inspection is seen as a cognitive hallmark purportedly indicating the existence of self-recognition. Only a few species of great apes have been reported to pass a standard mark test for mirror self-recognition in which animals attempt to touch a mark. In addition, evidence for passing the mark test was also reported for Asian elephants, two species of corvids, and a species of cleaner fish. Mirror self-recognition has also been claimed for bottlenose dolphins, using exposure of marked areas to a mirror as evidence. However, what counts as self-directed behaviour to see the mark and what does not has been debated. To avoid this problem, we marked the areas around both eyes of the animals at the same time, one with visible and the other with transparent dye to control for haptic cues. This allowed the animal to see the mark easily and us to investigate what side was exposed to the mirror as an indicator for mark observation. We found that the animals actively chose to inspect their visibly marked side while they did not show an increased interest in a marked conspecific in the pool. These results demonstrate that dolphins use the mirror to inspect their marks and, therefore, likely recognise a distinction between self and others.

Lateralization of visuospatial processing in the bottlenose dolphin (Tursiops truncatus).

Two adult female bottlenose dolphins were tested for cerebral asymmetries in the visuospatial domain. The animals learned under binocular conditions a three-choice spatial discrimination task with three hoops positioned along a line in the middle of the tank. During a correct trial the dolphins had to swim from a starting position at the tanks wall through one of the hoops, come back to the starting position, choose another hoop, swim back to start and finally swim through the third hoop. For such a trial to be correct, the animals had to swim through all three hoops in any sequence without omitting or re-using one of them. After reaching criterion binocularly, monocular trials (one eye covered with an adherent suction cup) were introduced where the dolphins carried out the same task alternatingly under left or right eye seeing conditions. For both animals, the right eye performance was clearly superior to that of the left eye. Binocular and right eye performances were similar. As a result of the complete decussation at the optic nerve, this right eye superiority suggests a left-hemispheric dominance for the processing of visuospatial information. This is a remarkable deviation from the usual right hemisphere advantage for these kind of tasks found in different species of mammals and birds.

Visual lateralization of pattern discrimination in the bottlenose dolphin (Tursiops truncatus).

The aim of the present study was to investigate whether bottlenose dolphins have cerebral asymmetries of visual processing. The monocular performance of the adult dolphin Goliath was tested using a large number of simultaneous multiple pattern discrimination tasks. The experiments revealed a clear right eye advantage in the acquisition and the retention of pattern discriminations as well as asymmetries in the interhemispheric transfer of visual information. As a result of a complete decussation at the optic nerve, this right eye superiority is probably related to a left hemisphere dominance in visual processing.

Unexpected discrimination strategy used by pigeons.

Pigeons were conditioned with a symbolic matching-to-sample paradigm. Six visual patterns in 16 configurations were presented in an operant chamber where reinforcement was delivered next to the correct keys. A test with novel configurations, planned to demonstrate associative transitivity between equivalent stimuli, revealed instead a different but consistent behavior. An explanation for this behavior was sought in further tests and in a detailed post-hoc analysis of the training configuration. The analysis revealed an overall location imbalance of the comparison stimuli in relation to the reinforcement sites. This apparently led to response side biases controlled by these lateral stimuli. The results of additional tests accorded with this hypothesis. The directing effect was unequivocal when both lateral stimuli biased a response in the same direction. When the directing effects by these stimuli were opposite, the animals only solved the task when they could resort to a rote learning of individual configurations. The characteristics of the rules that were identified are contrasted with those that have previously been thought to operate during conditional discrimination learning.

Category discrimination by pigeons using five polymorphous features.

Eight pigeons were trained to discriminate between sets of color photographs of natural scenes. The scenes differed along five two-valued dimensions (site, weather, camera distance, camera orientation, and camera height), and all combinations of the feature values were used. One value of each dimension was designated as positive, and slides containing three or more positive feature values were members of the positive stimulus set. Thus, each feature had an equal, low, correlation with reinforcement, and all features had zero correlations with each other. Seven of the 8 pigeons learned this discrimination, and their responding came under the control of all five features. Within the positive and negative stimulus sets, response rates were higher to stimuli that contained more positive feature values. Once discrimination had been achieved, reversal training was given using a subset of the slides. In this subset, only a single feature was correlated with reinforcement. All pigeons learned this reversal successfully and generalized it to additional photographs with the same feature content. After reversal, the original reinforcement contingencies were reinstated, and training was continued using all the slides except those that had been used in reversal. Reversal generalized to these slides to some extent. Analysis of the response rates to individual slides showed that, compared with prereversal training, only the feature that had been subjected to reversal contingencies showed a reversed correlation with response rate. The remaining features showed the same correlation with response rate as they had before reversal training. Thus, reversal on some members of a category following category discrimination training led to generalization to stimuli within the category that were not involved in the reversal, but not to features that were not reversed. It is therefore inappropriate to refer to the pigeons as learning a concept.

Visual memory lateralization in pigeons.

Previous experiments employing simple visual discrimination tasks have revealed a cerebral lateralization in the visual system of pigeons with a dominance of the left hemisphere. Until now, visual memory lateralization in birds has not been investigated. To study possible asymmetries of visual memory functions, a simultaneous instrumental discrimination procedure was used. The animals were trained to discriminate 100 different visual patterns from a further 625 similar stimuli. Retention tests were conducted under binocular and monocular conditions. When the subjects looked monocularly, retention performance was significantly higher with the right eye (left hemisphere) than with the left eye (right hemisphere). The results suggest that the lateralization of the pigeon's visual system depends at least partly on an asymmetry in visual memory capacity.