Learning of magnetic compass directions in pigeons

A proof of magnetic compass learning by pigeons under laboratory conditions has been attempted for decades, but all experiments have failed so far. The aim of the present study was to test whether pigeons can learn magnetic compass directions in an operant chamber if magnetic cues are presented as true spatial cues. Experimental sessions were carried out in the local geomagnetic field and in magnetic fields with matched total intensity and inclination, but different directions generated with Helmholtz-coils. Birds demonstrated successful learning with a performance level comparable to that in learning studies with magnetic anomalies. In addition, we compared the data from magnetic learning in the laboratory with performance from homing experiments in the field. The birds that were more successful in the learning experiment had vanishing bearings farther away from the home direction than the group mean at unfamiliar, but not at familiar sites. This might suggest that better learners explore unknown locations in a different way. Our findings represent the first evidence for operant magnetic compass learning in pigeons and also provide a link between behavioural data from the field and the laboratory.     

Different responses in two strains of chickens (<Emphasis Type="Italic">Gallus gallus</Emphasis>) in a magnetic orientation test

Previous studies demonstrated that layer strain domestic chicks bred for egg production can orient using directional cues from the magnetic field; here we report that chicks from a broiler strain bred for meat production do not use magnetic cues for orientation. We imprinted both strains of chicken on a red ball and subsequently trained them in a featureless testing arena. Between rewarded trials in the geomagnetic field, we inserted unrewarded tests under the following conditions: (1) in the geomagnetic field, (2) in a magnetic field with North shifted by 90 degrees and (3) in a magnetic field with the inclination inverted. The layer chicks made a correct axial response in 75-80% of the tests, shifting their choices following a rotation of magnetic North. Chicks of the broiler strain, in contrast, performed at chance level with between 47 and 60% of choices on the correct axis. This difference between the strains does not appear to be due to substantial strain differences in motivation to perform the task. It therefore appears possible that the selection of the broiler strain has led to the elimination of the specific ability to respond to magnetic cues in the test situation.     

Effects of ageing on visual discrimination learning in Drosophila melanogaster

Two experiments examined conditioned suppression of the Proboscis Extension Response (PER), unconditionally released by sucrose stimulation of gustatory tarsal receptors, in young (7-day-old), middle-aged (30-day-old) and old (50-day-old) Drosophila melanogaster males, reared at 25°C. Individual flies were trained in a differential conditioning procedure in which a white stimulus signalled a quinine reinforcer, whereas a black stimulus was non-reinforced. When trained from the outset with the discriminative procedure, flies of all ages acquired the discrimination, although the acquisition of PER suppression to the white stimulus was retarded in middle-aged and old flies. The retardation of the acquisition of PER suppression in middle-aged and old flies was replicated in a second study in which the flies received simple conditioning to the white stimulus prior to discrimination training.     

Comprehending compass directions

Two experiments investigated the processes by which compass directions are comprehended. In both experiments, the task was as follows. A compass direction (e.g., 210 deg) was visually presented. The subject drew a representation of the direction and then pressed a key. Reaction time from onset of the stimulus to the keypress was measured. The results suggested a model of direction comprehension involving two steps: first, computing the nearest cardinal direction to the target direction, and then “rotating” from the cardinal to the target direction. Rotation could be performed equally well clockwise or counterclockwise. Additionally, north-south. tended to be comprehended faster than east-west, confirming past results that have shown an advantage of up-down over left-right in discrimination tasks.     

An experimental study of strategy development

Three experiments are reported in which the strategy usage for a compass point directions task was investigated. For this task, people may use either an efficient cancellation strategy or an inefficient spatial strategy (see Roberts, Gilmore, & Wood, 1997). Experiment 1, together with Experiment 2, showed that users of cancellation are unlikely to have explicit prior knowledge of this strategy; instead they discover it as a result of experience with the task. Experiment 1 also showed that strategy selection is unlikely to be based on stylistic preferences, and that reducing the cognitive demands of the task does not increase the discovery of cancellation. Experiment 3 showed that evaluation of strategy validity is also of importance: Unsuccessful evaluation may prevent the selection of a newly discovered strategy, even if it is the more efficient method.     

Experiencing extinction within a task makes nonextinguished information learned within a different task context-dependent

In two experiments with rats, we analyzed the effect of experiencing extinction in one task on the context specificity of a new association learned within a different task. Rats were trained to run in a runway for water in Task 1, and received taste aversion conditioning in Task 2 (the tasks were reversed in Exp. 2). Half of the rats received conditioning and extinction of Task 1 in Context A, whereas the other half received no extinction. Then all animals received training in the alternate task in Context B, prior to testing in Context C. When they were tested in Context C, Task 2 performance was attenuated if Task 1 had been extinguished prior to Task 2. These results are similar to those we have reported in humans, and consistent with the idea that extinction prompts attention to contexts, regardless of whether or not the contexts were involved in extinction.     

Sleep deprivation selectively impairs memory consolidation for contextual fear conditioning

Many behavioral and electrophysiological studies in animals and humans have suggested that sleep and circadian rhythms influence memory consolidation. In rodents, hippocampus-dependent memory may be particularly sensitive to sleep deprivation after training, as spatial memory in the Morris water maze is impaired by rapid eye movement sleep deprivation following training. Spatial learning in the Morris water maze, however, requires multiple training trials and performance, as measured by time to reach the hidden platform is influenced by not only spatial learning but also procedural learning. To determine if sleep is important for the consolidation of a single-trial, hippocampus-dependent task, we sleep deprived animals for 0–5 and 5–10 h after training for contextual and cued fear conditioning. We found that sleep deprivation from 0–5 h after training for this task impaired memory consolidation for contextual fear conditioning whereas sleep deprivation from 5–10 h after training had no effect. Sleep deprivation at either time point had no effect on cued fear conditioning, a hippocampus-independent task. Previous studies have determined that memory consolidation for fear conditioning is impaired when protein kinase A and protein synthesis inhibitors are administered at the same time as when sleep deprivation is effective, suggesting that sleep deprivation may act by modifying these molecular mechanisms of memory storage.     

Simultaneous training on two hippocampus-dependent tasks facilitates acquisition of trace eyeblink conditioning

A common cellular alteration, reduced post-burst afterhyperpolarization (AHP) in CA1 neurons, is associated with acquisition of the hippocampus-dependent tasks trace eyeblink conditioning and the Morris water maze. As a similar increase in excitability is correlated with these two learning paradigms, we sought to determine the interactive behavioral effects of training animals on both tasks by using either a consecutive or simultaneous training design. In the consecutive design, animals were trained first on either the trace eyeblink conditioning task for six sessions, followed by training on the water maze task for six sessions, or vice versa. The simultaneous design consisted of six or 11 training days; animals received one session/day of both trace eyeblink conditioning and water maze training. Separate groups were used for consecutive and simultaneous training. Animals trained on both tasks simultaneously were significantly facilitated in their ability to acquire the trace eyeblink conditioning task; no effect of simultaneous training was seen on the water maze task. No effect was seen on acquisition for either task when using the consecutive training design. Taken together, these findings provide insight into how the hippocampus processes information when animals learn multiple hippocampus-dependent tasks.     

Discrimination of cardinal compass directions

The study tested the hypothesis that east-west is more difficult to discriminate than north-south because pre-experimentally the former is associated with right-left and the latter with up-down (or front-behind). Forty-eight undergraduates first learned north, south, east, and west in slides of unknown places. In one condition the compass directions were indicated by labels. Arrows presented in the plane of the slides were in other conditions used to create different associations between the compass directions and the egocentric directions right-left and front-behind. In a subsequent RT task subjects indicated if the directions were the same or different in two consecutively presented slides. No reliable effects were found for same responses but different responses supported the hypothesis in being slower for east-west than north-south when associated with right-left (or left-right) and faster when associated with front-behind (or behind-front).     

The Influence of Background Stimuli on Summation in Autoshaping

Five experiments examined the factors that determine whether or nor summation will occur when experimental stimuli are presented on a television screen for autoshaping with pigeons. In Experiment 1 there were periods when the television screen was illuminated white and periods when it was dark. When conditioning was conducted during the white periods, then summation was observed if the test trials were conducted with a white screen during the intertrial interval. Summation was not found when conditioning was conducted during the dark periods, no matter how testing was conducted. The remaining experiments demonstrated that summation will occur if at least some of the conditioning trials are conducted with the experimental stimuli being accompanied by white illumination in the remainder of the television screen. It was not necessary for this illumination to persist into the interval between trials. These findings indicate that summation is influenced by the similarity of the patterns of stimulation that are used for training and testing.     

Effects of post-training hippocampal injections of midazolam on fear conditioning

Benzodiazepines have been useful tools for investigating mechanisms underlying learning and memory. The present set of experiments investigates the role of hippocampal GABA(A)/benzodiazepine receptors in memory consolidation using Pavlovian fear conditioning. Rats were prepared with cannulae aimed at the dorsal hippocampus and trained with a series of white noise-shock pairings. In the first experiment, animals received intrahippocampal infusion of midazolam or vehicle immediately or 3 h after training. Then, 24 h later, freezing to the training context and the white noise were measured independently. Results show infusion of midazolam immediately, but not 3 h, after training selectively attenuates contextual fear conditioning. In the second experiment, animals received intrahippocampal infusions of an antisense oligodeoxynucleotide (ODN) targeting the alpha5 subunit of the GABA(A) receptor or a missense control for several days prior to training and testing. Immediately after training, animals received an infusion of either midazolam or vehicle. Western blots conducted after testing showed a significant decrease in alpha5-containing GABA(A) receptor protein. This reduction did not alter the effectiveness of midazolam immediately after training at impairing context fear memory. Therefore, alpha5-containing GABA(A) receptors may not contribute to the effects of midazolam on context fear conditioning when given immediately post-training.     

Differentiated ratings of perceived exertion during physical conditioning of older individuals using leg-weight loading.

Use of leg weights for physical conditioning was evaluated in 8 middle-aged male Ss; four Ss of similar age served as a control group. Pre- and post-training evaluation consisted of heart rate and oxygen uptake responses to five submaximal work loads which involved either level walking or cycling. Differentiated ratings of perceived exertion elicited for each work load were: a local muscular rating; a central or cardio-pulmonary rating; and an over-all or general rating. Submaximal heart rate decreased 6 to 9 beats/min. from pretraining values for all work load after training. The differentiated ratings for training generally reflected a reduced strain on the cardiovascular system and also improved functioning of the working muscles with training. However, when one set of sensations dominated the exertional perception the others appear to have been perceptually de-emphasized. Local muscular factors seemed to dominate the exertional perception for cycling, but central factors appeared to play a more important role for treadmill walking, at least within the range of velocities investigated.     

The effect of overtraining on transfer between tasks involving the same stimulus dimension

Two experiments are reported in which transfer effects between two discrimination tasks are examined following differing amounts of training on the first task. The two tasks were a conditional (successive) discrimination involving black/white cues, and a simultaneous brightness discrimination involving the same black/white cues.

In Experiment I the conditional task was presented first and in Experiment II the simultaneous task was presented first. The results of Experiment I show overall negative transfer which is not directly related to the amount of training. The results of Experiment II, however, reveal positive transfer although again the amount of training produced no significant effect. These seemingly discrepant findings are explained in terms of the difficulty of the conditional task and the development of rigid position stereotypes. It is argued that the concept of frustration-instigated behaviour is necessary to account for the results.     

The influence of temporal factors on automatic priming and conscious expectancy in a simple reaction time task

In a previous study, we reported a dissociation between subjective expectancy and motor behaviour in a simple associative learning task (Perruchet, Cleeremans, & Destrebecqz, 2006). According to previous conditioning studies (Clark, Manns, & Squire, 2001), this dissociation is observed when the to-be-associated events coterminate and thus overlap in time (a training regimen called delay conditioning), but not when they are separated by a temporal delay (trace conditioning). In this latter situation indeed, there tends to be a direct relationship between subjective expectancy and behaviour. In this study, we further investigated this issue in a series of experiments where conscious and unconscious components of performance were pitted against each other. In Experiments 1–3, participants performed a simple reaction time task in which a preparatory signal (a tone) either overlapped with or terminated earlier than the imperative stimulus (a visual target presented in 50% of the trials). After each response, participants also had to state how much they expected the imperative stimulus to be displayed on the next trial. Results indicate that reaction times tend to decrease when the tone is consistently followed by the visual target across successive trials, whereas conscious expectancy for the target decreases at the same time. Importantly, we systematically found that the temporal relationship between the tone and the target failed to influence performance. In a fourth experiment, we examined whether these results extend to a two-choice reaction time task. To our surprise, we observed a direct relationship between subjective expectancies and reaction time in that situation. We nevertheless observed that the introduction of a delay between the tone and the target had, once again, no effect on performance.     

Analysis of treatments to alleviate forgetting in rats

Two experiments with rats investigated the effectiveness of prior-cuing treatments for alleviating forgetting of aversive conditioning. The aim was to see which retrieval cues would be most effective within different contexts. Experiment 1 examined the contexts of classical fear conditioning and instrumental avoidance training. The results indicated that the response components were sufficient to reinstate avoidance training, whereas the unconditioned stimulus (US) was most effective for classical fear conditioning. In Experiment 1, the reinforcer per se was ineffective in reinstating instrumental avoidance training. Experiment 2 manipulated the training context and found that the US could be made an effective prior-cuing treatment for instrumental training if classical conditioning components were more prevalent during training. These results are interpreted to mean that a "critical context" must be reinstated by the cuing treatment if this treatment is to promote retrieval of the memory.     

Aversive conditioning under conditions of restricted awareness: Effects on spatial cueing

In the literature on aversive conditioning there is still debate on the role of awareness. According to some authors, affective learning can occur with or without contingency learning (dual-process model), whereas others argue that a single process produces both affective responses and contingency knowledge. Although many studies have investigated these models, the results to date are inconclusive. Based on a review of the literature, a new series of experiments was designed to examine aversive conditioning in the absence of contingency awareness. In the present study we examined the effects of subliminal aversive conditioning on a spatial cueing task. Awareness was stringently tested after conditioning. Three kinds of awareness were distinguished: contingency awareness (awareness of the CS?US contingencies, where CS is the conditioned stimulus, and US is the unconditioned stimulus), perceptual awareness (awareness of the perceptual differences between the CSs), and US expectancy (awareness of a threat feeling when confronted with the CS+, but not when confronted with the CS?). The results of three experiments demonstrated that responses on the spatial cueing task were modulated by subliminal aversive conditioning. Importantly, none of the participants was contingency aware or able to perceptually discriminate between the conditioned stimuli. However, in Experiment 3, only those participants showing some level of postconditioning expectancy awareness exhibited conditioning effects. These experiments suggest that subliminal aversive conditioning produces small but significant effects, which may be modulated by expectancy awareness.     

Cue competition between elementary trained stimuli: US miscuing, interference, and US omission

The present series of experiments was designed to assess whether rule-based accounts of Pavlovian learning can account for cue competition effects observed after elemental training. All experiments involved initial differential conditioning training with A-US and B alone presentations. Miscuing refers to the fact that responding to A is impaired after one B-US presentation whereas interference is the impairment of responding to A after presentation of C-US pairings. Omission refers to the effects on B of A alone presentations. Experiments 1-2a provided clear evidence for miscuing whereas interference was not found after 1, 5 or 10 C-US pairings. Moreover, Experiments 3 and 3a found only weak evidence for interference in an A-US, B ∣ C-US, D ∣ A design used previously to show the effect. Experiments 4 and 5 failed to find any effect of US omission after one or five omission trials. The present results indicate that miscuing is more robust than is the interference effect. Moreover, the asymmetrical effects of US miscuing and US omission are difficult to accommodate within rule-based accounts of Pavlovian conditioning.     

Cross-modal transfer of conditioned suppression in rats: Effects of US intensity and extinction of the initial conditioning task

In conditioned suppression of water licking behavior by rats, we obtained data indicating general transfer of fear conditioning. A series of experiments resulted in two major findings. First, pairing of a neutral stimulus with a shock in the initial conditioning task facilitated acquisition of subsequent fear conditioning to another neutral stimulus, if the conditioned fear of the initial task was extinguished prior to the second task and if equally strong shocks were employed in both conditioning tasks. Second, omission of the extinction treatment or employment of weaker shocks in the initial task resulted in retardation, rather than facilitation, of the second conditioning task. An application to human clinical settings is discussed.     

Conditioned craving cues elicit an automatic approach tendency

In two experiments, we used a Pavlovian differential conditioning procedure to induce craving for chocolate. As a result of repeated pairing with chocolate intake, initially neutral cues came to elicit an automatic approach tendency in a speeded stimulus-response compatibility reaction time task. This automatic approach tendency, moreover, seemed to be sensitive to manipulations of extinction and renewal in the Pavlovian conditioning procedure. These findings corroborate and extend previous reports of automatic approach tendencies elicited by substance-relevant cues in addiction, while controlling for alternative accounts for such observations. Moreover, our data lend support to and extend learning models of cue-induced craving and addiction. Finally, we argue that the procedure we present here provides an ecologically valid behavioural tool that allows studying processes involved in cue-induced craving, addiction and relapse without relying on verbal report.     

Deficits in trace cued fear conditioning in galanin-treated rats and galanin-overexpressing transgenic mice

Galanin inhibits the release of several neurotransmitters and produces performance deficits in a variety of spatial and aversive learning and memory tasks. The experiments in this study investigated the role galanin has in emotional learning and memory using a standard delay cued and contextual fear conditioning task. Rats were administered galanin into the lateral ventricles before training, and scored for freezing behavior in the same context and in a novel context with and without an auditory cue (CS) that had been paired previously with an aversive stimulus (US). Galanin-overexpressing transgenic mice were tested in an identical behavioral protocol. The galanin-administered rats and the transgenic mice were not significantly different from their respective controls on this task. A more challenging trace cued and contextual fear conditioning procedure was administered to separate groups of galanin-treated rats and galanin-overexpressing transgenic mice. Subjects were trained with the same CS and US, however, a 2.5-sec delay was inserted between CS offset and US onset. Following the trace conditioning, rats administered galanin and mice overexpressing galanin both exhibited significantly less freezing to the CS in the novel context as compared with their control groups. These results indicate that the observed disruption of cued fear conditioning was specific to the more difficult trace conditioning task. These findings are the first demonstration that galanin impairs performance on an emotional memory task and support the hypothesis that galanin-induced deficits are specific to more difficult cognitive tasks.