Bees were incrementally trained to enter the Y-maze and both arms of the apparatus over 30- to 60-min periods. Once each bee was able to fly into the entrance hole and the hole that led to the decision chamber and could find the poles in both Y-maze arms, the experiment began.

After entering the Y-maze, bees would be in the initial chamber where they could view the sample number. To solve the task, the bees were required to either add or subtract the value of one to this sample number depending on the color of the elements (Fig. 1). Bees would then fly through the next hole in the Y-maze and into the decision chamber where they could simultaneously view two stimuli in a dual choice test. If the sample number was blue, then the bee would need to choose the option that was one element greater than the sample stimulus to receive a reward, while if the sample number was yellow, then the bee would need to choose the option that was one element less than the sample number to receive a reward. The incorrect option was randomly selected and could be any number from 1 to 5, including the sample number itself that controlled for the bees choosing the correct option based on visual similarity, and incorrect choices were associated with a bitter tasting quinine solution.

Each bee thus completed 100 appetitive-aversive (31) reinforced trials presenting either addition or subtraction arithmetic problems. Whether a trial would involve adding or subtracting one element from the sample number was randomized.

Throughout the training, the numbers that could be used for the sample in the addition trials were 1, 2, and 4. Thus, correct answers could be 2, 3, and 5 and the incorrect answers could be 1, 2, 3, 4, and 5. During the subtraction trials, the numbers that could be used for the sample number were 2, 4, and 5. Thus, correct answers could be 1, 3, and 4 and the incorrect answers could be 1, 2, 3, 4, and 5. The number 3 was never shown as a sample number during training for any bee and was thus used as the sample number for all unreinforced tests to ensure the sample number was novel during tests.

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