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We studied the ability of honeybees to discriminate between single odorants and binary olfactory mixtures. We analyzed the effect of the number of common elements between these two stimulus classes on olfactory discrimination. We used olfactory conditioning of the honeybees' proboscis extension reflex (PER), a paradigm in which odors can be associated with a reinforcement of sucrose solution. Bees were asked to discriminate reinforced from nonreinforced olfactory stimuli. They were trained with two elemental odors (A and B) versus a binary olfactory mixture. The mixture was either AB (group 2CE, two common elements), AC (group 1CE, one common element A), or CD (group 0CE, no common element). Three groups followed a positive patterning schedule (mixture reinforced and elements nonreinforced: groups 2CE+, 1CE+, and 0CE+) and three other groups a negative patterning schedule (mixture nonreinforced and elements reinforced: groups 2CE−, 1CE−, and 0CE−). We showed that a reduction of similarity (number of common elements) between elemental odors and compounds enhanced the ability to discriminate elements from compounds and that the kind of compound processing used by the bees supports theories that assume nonelemental compound processing (i.e., that exclude the mere summation of the elemental associative strengths upon compound presentation).
We studied the ability of honeybees to discriminate between single odorants and binary olfactory mixtures. We analyzed the effect of the number of common elements between these two stimulus classes on olfactory discrimination. We used olfactory conditioning of the honeybees' proboscis extension reflex (PER), a paradigm in which odors can be associated with a reinforcement of sucrose solution. Bees were asked to discriminate reinforced from nonreinforced olfactory stimuli. They were trained with two elemental odors (A and B) versus a binary olfactory mixture. The mixture was either AB (group 2CE, two common elements), AC (group 1CE, one common element A), or CD (group 0CE, no common element). Three groups followed a positive patterning schedule (mixture reinforced and elements nonreinforced: groups 2CE+, 1CE+, and 0CE+) and three other groups a negative patterning schedule (mixture nonreinforced and elements reinforced: groups 2CE−, 1CE−, and 0CE−). We showed that a reduction of similarity (number of common elements) between elemental odors and compounds enhanced the ability to discriminate elements from compounds and that the kind of compound processing used by the bees supports theories that assume nonelemental compound processing (i.e., that exclude the mere summation of the elemental associative strengths upon compound presentation).
We investigated the capability of honeybees to discriminate between single odorants, binary olfactory mixtures, and ternary olfactory mixtures in olfactory conditioning of the proboscis extension reflex. In Experiment 1, three single odorants (A+, B+, and C+) and three binary mixtures of these odors (AB+, AC+, and BC+) were reinforced while the ternary compound, consisting of all three odors (ABC−), was nonreinforced. In Experiment 2, only one single odorant (A+) and one binary olfactory compound (BC+) were reinforced while the ternary compound (ABC−) consisting of the single odor and the binary compound was nonreinforced. We studied whether bees can solve these problems and whether the course of differentiation can be predicted by the unique cue theory, a modified unique cue theory, or Pearce's configural theory. Honeybees were not able to differentiate reinforced from nonreinforced stimuli in Experiment 1. However, summation to ABC observed at the beginning of training contradicts the predictions of Pearce's configural theory. In Experiment 2, differentiation between the single odorant A and the ternary compound developed more easily than between the binary compound BC and ABC. This pattern of differentiation is in line with a modified unique cue theory and Pearce's configural theory. Summation to ABC at the beginning of training, however, again was at odds with Pearce's configural theory. Thus, olfactory compound processing in honeybees can best be explained by a modified unique cue theory.The processing of stimuli composed of more than one single element (compounds) is of major interest but has not been clearly elucidated up to now (e.g., Rowe 1999; Pearce and Bouton 2001). Two main approaches, an "elemental" and a "configural" one, have been proposed to explain compound processing. The former assumes that animals are able to extract the elemental composition of the compound, whereas the latter postulates that animals process a compound as a new configuration, independently of its single stimuli. Among elemental theories, the pure elemental theory is the most simple and supposes that the total associative strength (V) of a compound AB results from the mere summation of the associative strengths of its elements A and B (i.e., V AB = V A + V B ). For each stimulus, the change in associative strength, ⌬V, results from the model of Rescorla and Wagner (1972):where ⌬V is the change in associative strength V, ␣ is the learning rate associated with the stimulus under consideration (A, B, or AB),  is the learning rate associated with the reinforcement, is the asymptotic level of associative strength that can be supported by the reinforcement, and V T is the combined associative strengths of all stimuli present on a learning trial. An extension of the elemental theory is the unique cue theory, which assumes that a compound AB is processed as the sum of the single elements A and B, plus a stimulus U, which is unique to the compound and results from the conjunction of A and B (i.e., Rescorla 1972Rescorla , 1973Whitlow...
Theories of associative learning are concerned with the factors that govern association formation when two stimuli are presented together. In this article we review the relative merits of the currently influential theories of associative learning. Some theories focus on the role of attention in association formation, but differ in the rules they propose for determining whether or not attention is paid to a stimulus. Other theories focus on the nature of the association that is formed, but differ as to whether this association is regarded as elemental, configural, or hierarchical. Recent developments involve modifications to existing theories in order to account for associative learning between two stimuli, A and B, when A is accompanied, not by B, but by a stimulus that has been paired with B. The implications of the theories for understanding how humans derive causal judgments and solve categorization problems is considered.
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