Reversal of a Spatial Discrimination Task in the Common Octopus (Octopus vulgaris).

Reversal learning requires an animal to learn to discriminate between two stimuli but reverse its responses to these stimuli every time it has reached a learning criterion. Thus, different from pure discrimination experiments, reversal learning experiments require the animal to respond to stimuli flexibly, and the reversal learning performance can be taken as an illustration of the animal's cognitive abilities. We herein describe a reversal learning experiment involving a simple spatial discrimination task, choosing the right or left side, with octopus. When trained with positive reinforcement alone, most octopuses did not even learn the original task. The learning behavior changed drastically when incorrect choices were indicated by a visual signal: the octopuses learned the task within a few sessions and completed several reversals thereby decreasing the number of errors needed to complete a reversal successively. A group of octopus trained with the incorrect-choice signal directly acquired the task quickly and reduced their performances over reversals. Our results indicate that octopuses are able to perform successfully in a reversal experiment based on a spatial discrimination showing progressive improvement, however, without reaching the ultimate performance. Thus, depending on the experimental context, octopus can show behavioral flexibility in a reversal learning task, which goes beyond mere discrimination learning.

Reconsideration of Serial Visual Reversal Learning in Octopus (Octopus vulgaris) from a Methodological Perspective.

Octopuses (Octopus vulgaris) are generally considered to possess extraordinary cognitive abilities including the ability to successfully perform in a serial reversal learning task. During reversal learning, an animal is presented with a discrimination problem and after reaching a learning criterion, the signs of the stimuli are reversed: the former positive becomes the negative stimulus and vice versa. If an animal improves its performance over reversals, it is ascribed advanced cognitive abilities. Reversal learning has been tested in octopus in a number of studies. However, the experimental procedures adopted in these studies involved pre-training on the new positive stimulus after a reversal, strong negative reinforcement or might have enabled secondary cueing by the experimenter. These procedures could have all affected the outcome of reversal learning. Thus, in this study, serial visual reversal learning was revisited in octopus. We trained four common octopuses (O. vulgaris) to discriminate between 2-dimensional stimuli presented on a monitor in a simultaneous visual discrimination task and reversed the signs of the stimuli each time the animals reached the learning criterion of ≥80% in two consecutive sessions. The animals were trained using operant conditioning techniques including a secondary reinforcer, a rod that was pushed up and down the feeding tube, which signaled the correctness of a response and preceded the subsequent primary reinforcement of food. The experimental protocol did not involve negative reinforcement. One animal completed four reversals and showed progressive improvement, i.e., it decreased its errors to criterion the more reversals it experienced. This animal developed a generalized response strategy. In contrast, another animal completed only one reversal, whereas two animals did not learn to reverse during the first reversal. In conclusion, some octopus individuals can learn to reverse in a visual task demonstrating behavioral flexibility even with a refined methodology.