Second-Order Olfactory and Visual Learning in the Optic Tectum of the Goldfish

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Abstract
1. The goldfish (Carassius auratus) learns to swim to concealed food when an illuminated disk is presented. 2. The presentation of an olfactory stimulus at the same time as the optic stimulus, during reinforcement of this visual learning, results in the addition of the olfactory stimulus to the stimulus complex necessary to call forth the learned reaction. 3. Five animals learned to react to a situation in which--after preliminary training with an optic stimulus with food as reward--amyl acetate was given as olfactory stimulus and the optic stimulus as reward. This effect is termed second-order learning. It is comparable to second-order conditioned reflexes. 4. In four animals trained in this way operations involving removal of large areas of the optic tectum, or cuts made at its anterior border, caused disturbances in the second-order learning. 5. There exists, therefore, in the Teleostean optic tectum a mechanism capable of second-order olfactory-optic learning. 6. The connexion between the telencephalic primary olfactory centre and the tectum involved in this learning passes into the tectum at its anterior border, since cuts made in this region interfered with the learning while leaving a great deal of the tectum intact.

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