Harald Lachnit

Neural representation of olfactory mixtures in the honeybee antennal lobe

Natural olfactory stimuli occur as mixtures of many single odors. We studied whether the representation of a mixture in the brain retains single‐odor information and how much mixture‐specific information it includes. To understand mixture representation in the honeybee brain, we used in vivo calcium imaging at the level of the antennal lobe, and systematically measured odor‐evoked activity in 24 identified glomeruli in response to four single odorants and all their possible binary, ternary and quaternary mixtures. Qualitatively, mixture‐induced activity patterns always contained glomeruli belonging to the pattern of at least one of the components, suggesting a high conservation of component information in olfactory mixtures. Quantitatively, glomerular activity saturated quickly and increasing the number of components resulted in an increase of cases in which the response of a glomerulus to the mixture was lower than that to the strongest component (‘suppression’). This shows global inhibition in the antennal lobe, probably acting as overall gain control. Single components were not equally salient (in terms of number of active glomeruli) and mixture activity patterns were always more similar to the more salient components, in a way that could be predicted linearly. Thus, although a gain control system in the honeybee antennal lobe prevents saturation of the olfactory system, mixture representation follows essentially elemental rules.

Positive and negative patterning in human classical skin conductance response conditioning

Three experiments on classical differential conditioning of the human skin conductance response to elemental and compound stimuli are reported. Subjects in Experiment 1 received both positive and negative patterning training, followed by either positive or negative patterning transfer tests on new stimuli. In positive patterning, a compound of two stimuli is reinforced and its elements are nonreinforced. In negative patterning, the elements are reinforced and the compound is nonreinforced. Subjects in Experiments 2 and 3 received either positive or negative patterning during training, followed by transfer tests on new stimuli. In Experiment 2, the transfer series began with new elements, after which their compound was presented; in Experiment 3, the new compound was presented first in the transfer series, and then the separate elements were administered. All three experiments provided evidence of the acquisition of positive patterning, while negative patterning was found only in Experiments 2 and 3. Positive patterning transferred to new stimuli, indicating that it was not attributable solely to summation of sub-threshold excitation conditioned to the elements on reinforced compound trials. This finding, coupled with the negative patterning found in Experiments 2 and 3, provided support for the unique cue hypothesis. It was concluded that the assumed unique cue constituted a learned “rule,” and that the actual elemental stimuli were neither perceptually nor otherwise modified during the conditioning process.

Stimulus coding in human associative learning: Flexible representations of parts and wholes

An enduring theme for theories of associative learning is the problem of explaining how configural discriminations--ones in which the significance of combinations of cues is inconsistent with the significance of the individual cues themselves-are learned. One approach has been to assume that configurations are the basic representational form on which associative processes operate, another has tried in contrast to retain elementalism. We review evidence that human learning is representationally flexible in a way that challenges both configural and elemental theories. We describe research showing that task demands, prior experience, instructions, and stimulus properties all influence whether a particular problem is solved configurally or elementally. Lines of possible future theory development are discussed.

Within-compound associations in retrospective revaluation and in direct learning: A challenge for comparator theory

In three human causal learning experiments we investigated the role of within-compound associations in learning about absent cues versus learning about present cues. Different theoretical approaches agree that within-compound associations are essential for learning about absent cues-that is, for retrospective revaluation. They differ, however, with regard to the role of within-compound associations for learning about present cues-that is, for direct learning. A memory test was used to assess within-compound associations. Experiment 1 used a blocking/release from overshadowing design, Experiment 2 used a conditioned inhibition design, and Experiment 3 used a higher-order cue selection design. In all experiments, first-order retrospective revaluation was significantly correlated with within-compound associations, but no significant correlations were found for the direct learning conditions. In addition to this, second-order retrospective revaluation in Experiment 3 was positively correlated to joint knowledge of first-order and second-order within-compound associations. Furthermore, cue selection effects were stronger for direct learning conditions than for retrospective learning conditions. These results are at variance with the comparator hypothesis but are in agreement with a modified associative theory and with the suggestion that retrospective revaluation might be due to rehearsal processes.

The effect of cumulative experience on the use of elemental and configural visual discrimination strategies in honeybees

We addressed the question of whether the amount of individual experience determines the use of elemental or configural visual discrimination strategies in free-flying honeybees Apis mellifera. We trained bees to fly into a Y-maze to collect sucrose solution on a rewarded stimulus presented in one of the arms of the maze. Stimuli were colour disks, violet (V), green (G) or yellow (Y), which were of equal psychophysical salience for honeybees. Training followed an A+, BC+ design, followed by an AC versus BC test. Training consisted of 6 (3 A+ and 3 BC+), 20 (10 A+ and 10 BC+) or 40 (20 A+ and 20 BC+) acquisition trials. Elemental models of compound processing predict a preference for the non-trained stimulus AC while configural models predict a preference for the trained stimulus BC. Our results show that increasing the number of acquisition trials results in a change of the internal representation of stimuli. After six training trials, bees favoured an elemental strategy and preferred AC to BC during the tests. Generally, increasing the number of training trials resulted in an increase of the choice of BC. Thus, short training favoured processing of the compound as the sum of its elements (elemental account) while long training favoured its processing as being different from the sum of its elements (configural account). Additionally, we observed that the change in stimulus processing was also influenced by stimulus similarity. Colour perceptual similarity favoured configural processing with increasing experience.

Nonelemental visual learning in honeybees

Abstract Free-flying honeybees, Apis mellifera, learn visual stimuli in the appetitive context of food search. Visual compound stimuli are relevant in nautre as bees learn flower images that consist of many visual elements. We studied whether elemental associations between each visual element and the reinforcement (elemental approach) are enough to explain the solving of visual discrimination problems that raise ambiguity at the elemental level. We asked whether bees could solve three different visual discriminations: (1) positive patterning (A−, B−, AB+); (2) negative patterning (A+, B+, AB−); and (3) biconditional discrimination (AB+, CD+, AC−, BD−). In experiments 1 and 2 bees had to discriminate a yellow–violet chequerboard from the yellow or the violet squares alone. In experiment 3, four different gratings combining one colour (yellow or violet) with one orientation (vertical or horizontal) had to be discriminated. In all three problems binary compounds were trained in such a way that each element appeared equally often as rewarded and nonrewarded. Bees could solve the three discrimination problems. They always chose the reinforced stimulus despite ambiguity at the level of the elements. For solving positive patterning, elemental processing could be used. For negative patterning and biconditional discrimination, nonelemental processing strategies (unique-cue or configural approach) are necessary to account for these results. Although we cannot decide between a configural and a unique-cue interpretation, we can clearly reject purely elemental processing in these cases. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.

Non-elemental processing in olfactory discrimination tasks needs bilateral input in honeybees

In patterning discriminations, animals have to differentiate a compound stimulus AB from each of its elements A and B. In positive patterning (PP), the compound is reinforced whilst the single elements are non-reinforced. In negative patterning (NP), single elements are reinforced whilst the compound is non-reinforced. Using olfactory conditioning of the proboscis extension response (PER), we asked whether honeybees (Apis mellifera) can solve these patterning problems when odorants are given unilaterally as well as bilaterally to the antennae. Separating the olfactory input space of bees into two independent zones using plastic walls placed between the antennae, we conditioned bees in PP and NP procedures, with input on one side, on both sides, or in an ambiguous problem where bees had to solve PP on one side and NP on the other side. We found that bees with simultaneous bilateral input solve both patterning tasks efficiently. In contrast, PP but not NP was learned by bees receiving unilateral olfactory input. Bees subjected to the ambiguous NP/PP problem only solved PP. As PP can be solved through mere elemental processes, but NP is critically dependent on the use of non-elemental learning processes, our results suggest that bilateral olfactory input is necessary for non-elemental processing to take place in the bee brain.

Similarity and discrimination in human Pavlovian conditioning

We report three Pavlovian eyelid conditioning experiments with humans, designed to experimentally decide between elemental and configural learning theories. We used two different designs originally proposed by Redhead and Pearce (1995). In Experiments 1 and 2, three stimulus elements, A, B, and C, were presented in all possible combinations. All patterns were reinforced except for pattern ABC (A/B/C+, AB/AC/BC+, ABC-). According to elemental learning theories, response proportions on A/B/C+ trials should be smaller than on AB/AC/BC+ trials, whereas configural learning theory makes the opposite prediction. The results confirmed neither prediction. In Experiment 3, the A/B/C+, AB/AC/BC+, and ABC- trials were interspersed by D/E/F-, DE/DF/EF-, DEF+ trials. Again, neither prediction was confirmed. We suggest a modification of configural learning theory as a possible explanation of our results.

Contextual control in discrimination reversal learning.

In 3 human predictive learning experiments, the authors examined contextual control of responding in discrimination reversal learning. In Phase 1, a discrimination between 2 stimuli (A+, B-) was trained in Context 1. During Phase 2, participants received discrimination reversal training (A-, B+) in Context 2. Testing occurred in Context 1 and Context 2 (Experiments 1A and 1B) or in Context 1 and Context 3 (Experiment 2). During the test phase, performance in Context 1 and Context 2 reflected the contingencies trained during Phase 1 and Phase 2, respectively. When testing occurred in Context 3, there was no discriminative responding between A and B. In addition, the experiments demonstrated that discriminating stimuli with a consistent reinforcement history were also affected by contextual manipulations. Results indicate that each training context acquires the ability to control performance. Unique-cue and configural approaches account for a major part of the results.

Mobilization of Cognitive Resources and the Generation Effect

The generation effect refers to the memory advantage of words that have been generated rather than read. Such a read-generate comparison confounds qualitative task differences and raises methodological problems. A revised methodology is proposed circumventing these problems in that the encoding task is held constant and all stimuli have to be generated, but the degree of generativeness (i.e. the amount of cueing) is varied. In Experiment 1, 1, the (refined version of the) generation effect is demonstrated in a within-subjects design; with increasing generation activity left to the subject, free recall performance increases. No effect is obtained for degree of target masking. The same finding is replicated and shown to be independent of self-paced study time when generative activity is manipulated between subjects (Experiment 2) or within subjects (Experiment 3). As all learning trials involve generation, encoding time is controlled statistically, and free recall is used as a measure of memory, this refined generation effect cannot be explained as an artifact of selective attention or elaboration. Rather, generative activity seems to increase the mobilization of cognitive resources. This motivational account is supported by Experiment 4 showing an enhanced generation effect for positive mood.