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In an appetitive context, honeybees (Apis mellifera) learn to associate odors with a reward of sucrose solution. If an odor is presented immediately before the sucrose, an elemental association is formed that enables the odor to release the proboscis extension response (PER). Olfactory conditioning of PER was used to study whether, beyond elemental associations, honeybees are able to process configural associations. Bees were trained in a positive and anegative patterning discrimination problem. In the first problem, single odorants were nonreinforced whereas the compound was reinforced. In the second problem, single odorants were reinforced whereas the compound was nonreinforced. We studied whether bees can solve these problems and whether the ratio between the number of presentations of the reinforced stimuli and the number of presentations of the nonreinforced stimuli affects discrimination. Honeybees differentiated reinforced and nonreinforced stimuli in positive and negative patterning discriminations. They thus can process configural associations. The variation of the ratio of reinforced to nonreinforced stimuli modulated the amount of differentiation. The assignment of singular codes to complex odor blends could be implemented at the neural level: When bees are stimulated with odor mixtures, the activation patterns evoked at the primary olfactory neuropile, the antennal lobe, may be combinations of the single odorant responses that are not necessarily fully additive.
In an appetitive context, honeybees (Apis mellifera) learn to associate odors with a reward of sucrose solution. If an odor is presented immediately before the sucrose, an elemental association is formed that enables the odor to release the proboscis extension response (PER). Olfactory conditioning of PER was used to study whether, beyond elemental associations, honeybees are able to process configural associations. Bees were trained in a positive and anegative patterning discrimination problem. In the first problem, single odorants were nonreinforced whereas the compound was reinforced. In the second problem, single odorants were reinforced whereas the compound was nonreinforced. We studied whether bees can solve these problems and whether the ratio between the number of presentations of the reinforced stimuli and the number of presentations of the nonreinforced stimuli affects discrimination. Honeybees differentiated reinforced and nonreinforced stimuli in positive and negative patterning discriminations. They thus can process configural associations. The variation of the ratio of reinforced to nonreinforced stimuli modulated the amount of differentiation. The assignment of singular codes to complex odor blends could be implemented at the neural level: When bees are stimulated with odor mixtures, the activation patterns evoked at the primary olfactory neuropile, the antennal lobe, may be combinations of the single odorant responses that are not necessarily fully additive.
Rabbits were given reinforced training of the nictitating membrane (NM) response using separate conditioned stimuli (CSs), which were a tone, light, and/or tactile vibration. Then, two CSs were compounded and given further pairings with the unconditioned stimulus (US). Evidence of both overexpectation and summation effects appeared. That is, responding to the individual CSs declined despite their continued pairing with the US on compound trials (overexpectation), and responding on the compound trials was greater than responding to the individual CSs (summation). The response loss appeared regardless of the testing regime, that is, whether the test presentations of the individual CSs were themselves reinforced (Experiment 2), not reinforced (Experiment 1), or deferred until the end of compound training (Experiment 2). The results are discussed with respect to the roles of excitatory versus inhibitory processes, elemental versus configural processes, and the possible roles of cerebellar and hippocampal pathways.
The scientific study of associative learning began nearly 100 years ago with the pioneering studies of Thorndike and Pavlov, and it continues today as an active area of research and theory. Associative learning should be the foundation for our understanding of other forms of behavior and cognition in human and nonhuman animals. The laws of associative learning are complex, and many modern theorists posit the involvement of attention, memory, and information processing in such basic conditioning phenomena as overshadowing and blocking, and the effects of stimulus preexposure on later conditioning. An unresolved problem for learning theory is distinguishing the formation of associations from their behavioral expression. This and other problems will occupy future generations of behavioral scientists interested in the experimental investigation of associative learning. Neuroscientists and cognitive scientists will both contribute to and benefit from that effort in the next 100 years of inquiry.
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