Memories in drosophila heat-box learning

Learning and memory processes of operant conditioning in the heat-box are analyzed. In a search for conditioning parameters leading to high retention scores, intermittent training is shown to give better results than those of continuous training. Immediate retention tests contain two memory components, a spatial preference for one side of the chamber and a “stay-where-you-are-effect.” Intermittent training strengthens the latter. In the second part, memory dynamics is investigated. Flies are trained in one chamber and tested in a second one after a brief reminder training. With this direct transfer, memory scores reflect an associative learning process in the first chamber. To investigate memory retention after extended time periods, indirect transfer experiments are performed. The fly is transferred to a different environment between training and test phases. With this procedure, an aftereffect of the training can still be observed 2 h later. Surprisingly, exposure to the chamber without conditioning also leads to a memory effect in the indirect transfer experiment. This exposure effect reveals a dispositional change that facilitates operant learning during the reminder training. The various memory effects are independent of the mushroom bodies.     

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.     

Spatial contiguity facilitates Pavlovian second-order conditioning

Two experiments investigated the effects of spatial contiguity upon the formation of second-order conditioning in pigeon subjects. Experiment 1 used an autoshaping procedure to pair two visual stimuli, S2 and S1, after S1 had previously been paired with food. The resulting second-order conditioning of S2 was superior when both stimuli appeared on the same response key within a trial, compared with their appearing on different keys. Experiment 2 found a similar importance of spatial contiguity between S2 and S1 in a conditioned suppression paradigm. In addition, consistently presenting S2 and S1 in the same spatial location produced superior conditioning compared with varying their spatial relation from trial to trial. The design of these experiments was such as to imply that spatial contiguity facilitates performance by improving the formation of association rather than by promoting stimulus generalization or pseudoconditioning. Moreover, the observation of a facilitative effect of spatial contiguity between S1 and S2 in two different paradigms that use qualitatively different unconditioned stimuli and evoke different responses implies some generality for these findings. Consequently, these results suggest that spatially contiguous stimuli are especially associable in Pavlovian conditioning paradigms.     

Stimulus-specific sequence representation in serial reaction time tasks

Some recent evidence has favoured purely response-based implicit representation of sequences in serial reaction time tasks. Three experiments were conducted using serial reaction time tasks featuring four spatial stimuli mapped in categories to two responses. Deviant items from the expected sequence that required the expected response resulted in increased response latencies. The findings demonstrated a stimulus-specific form of representation that operates in the serial reaction time task. No evidence was found to suggest that the stimulus-specific learning was contingent on explicit knowledge of the sequence. Such stimulus-based learning would be congruent with a shortcut within an information-processing framework and, combined with other research findings, suggests that there are multiple loci for learning effects.     

Pavlovian eyeblink conditioning in combat veterans with and without post-traumatic stress disorder

Several recent studies have investigated relationships between post-traumatic stress disorder (PTSD) and learning and memory problems. These reports have found in general that not only does PTSD affect trauma-related memories, but when patients with PTSD are compared with similar trauma patients without PTSD, general memory impairments have been found. The present paper reports a study in which associative learning, using Pavlovian eyeblink conditioning, was investigated in combat veterans with and without chronic PTSD, using interstimulus intervals of 500 and 1000 msec in two separate experiments. Although several recent reports suggest that larger-magnitude autonomic conditioned responses occur in patients with PTSD during Pavlovian conditoning, the present study found evidence of impaired Pavlovian eyeblink conditioning in combat veterans with and without PTSD, compared to non-combat veterans. Although these data suggest that combat leads to an impaired associative learning process regardless of whether PTSD is apparent, a group of community-dwelling combat veterans not under medical treatment showed normal conditioning, suggesting that variables other than prior combat must also be involved.     

Pavlovian Eyeblink Conditioning in Combat Veterans With and Without Post-Traumatic Stress Disorder

Several recent studies have investigated relationships between post-traumatic stress disorder (PTSD) and learning and memory problems. These reports have found in general that not only does PTSD affect trauma-related memories, but when patients with PTSD are compared with similar trauma patients without PTSD, general memory impairments have been found. The present paper reports a study in which associative learning, using Pavlovian eyeblink conditioning, was investigated in combat veterans with and without chronic PTSD, using interstimulus intervals of 500 and 1000 msec in two separate experiments. Although several recent reports suggest that larger-magnitude autonomic conditioned responses occur in patients with PTSD during Pavlovian conditioning, the present study found evidence of impaired Pavlovian eyeblink conditioning in combat veterans with and without PTSD, compared to non-combat veterans. Although these data suggest that combat leads to an impaired associative learning process regardless of whether PTSD is apparent, a group of community-dwelling combat veterans not under medical treatment showed normal conditioning, suggesting that variables other than prior combat must also be involved.     

Trace eyeblink conditioning requires the hippocampus but not autophosphorylation of alphaCaMKII in mice

Little is known about signaling mechanisms underlying temporal associative learning. Here, we show that mice with a targeted point mutation that prevents autophosphorylation of alphaCaMKII (alphaCaMKII(T286A)) learn trace eyeblink conditioning normally. This forms a sharp contrast to the severely impaired spatial learning in the water maze and contextual fear conditioning observed in alphaCaMKII(T286A) mutants. Importantly, hippocampal lesions impaired trace eyeblink conditioning in alphaCaMKII(T286A) mice, suggesting a potential role of hippocampal alphaCaMKII-independent mechanisms. These results indicate that hippocampal signaling mechanisms that underlie temporal associative learning as assessed by trace eyeblink conditioning may differ from those of spatial and contextual learning.     

When goals constrain: Eye movements and memory for goal‐Oriented map study

Perspective goals, such as studying a map to learn a route through an environment or the overall layout of an environment, produce memory congruent with the goal‐directed rather than the studied perspective. One explanation for this finding is that perspective goals guide attention towards actively gathering relevant information during learning. A second explanation is that information is automatically organized into a goal‐congruent spatial model that guides retrieval. Both explanations predict goal‐congruent memory, but only the former one predicts eye movement differences during study. The present experiment investigated the effect of perspective goals on eye movement during map study and the flexibility of resulting spatial memories. Results demonstrate eye movements towards goal‐congruent map elements during learning, and lasting memory effects at test. These findings carry implications for the design of adaptive hand‐held and in‐vehicle navigation interfaces that accommodate for varied user goals. Copyright © 2008 John Wiley & Sons, Ltd.     

Evaluation and development of a connectionist theory of configural learning

A configural theory of associative learning is described that is based on the assumption that conditioning results in associations between the unconditioned stimulus and a representation of the entire pattern of stimulation that was present prior to its delivery. Configural theory was formulated originally to account for generalization and discrimination in Pavlovian conditioning. The first part of the article demonstrates how this theory can be used to explain results from studies of overshadowing, blocking, summation, and discrimination learning. The second part of the article shows how the theory can be developed to explain a broader range of phenomena, including mediated conditioning, reinforcer devaluation effects, the differential outcomes effect, acquired equivalence, sensory preconditioning, and structural discriminations.     

Associative change in the representations acquired during conditional discriminations: further analysis of the nature of conditional learning

Three experiments with rats investigated how the associative strengths of the representations that underlie conditional learning change when they are conditioned in compound. The results of each experiment suggest that the representation whose associative strength is most discrepant from the asymptote supported by the outcome of the trial undergoes the greatest change in associative strength. These results parallel those from simple Pavlovian conditioning (e.g., R. A. Rescorla, 2000). are inconsistent with unique-cue and configural accounts of conditional learning, and support a connectionist analysis of learning in which a "winner-takes-all" rule applies to the hidden units that can be activated and acquire associative strength at a given point in time.     

Selection history: How reward modulates selectivity of visual attention

Visual attention enables us to selectively prioritize or suppress information in the environment. Prominent models concerned with the control of visual attention differentiate between goal-directed, top-down and stimulus-driven, bottom-up control, with the former determined by current selection goals and the latter determined by physical salience. In the current review, we discuss recent studies that demonstrate that attentional selection does not need to be the result of top-down or bottom-up processing but, instead, is often driven by lingering biases due to the “history” of former attention deployments. This review mainly focuses on reward-based history effects; yet other types of history effects such as (intertrial) priming, statistical learning and affective conditioning are also discussed. We argue that evidence from behavioral, eye-movement and neuroimaging studies supports the idea that selection history modulates the topographical landscape of spatial “priority” maps, such that attention is biased toward locations having the highest activation on this map.     

Displacement limit (dmax) of sampled directional motion: direct and indirect estimates

The maximum displacement at which directional motion can be seen, known as dmax, has been said to define the spatial limits of the short-range motion system. Turano and Pantle (1985) used duration of motion aftereffect (MAE) to estimate the spatial limit of the short-range system, the assumption that dmax (a direct measure of motion perception) and MAE (an indirect measure) are equivalent indices of the same underlying perceptual process. In a series of four experiments, we examined this assumption by measuring dmax and duration of MAE across a range of displacements, stimulus waveforms (sine- or square-wave gratings), and spatial frequencies. We found that dmax and duration of MAE were affected differently by changes in the same variables. Therefore, we concluded that the two indices cannot be regarded as equivalent measures of the spatial limits of the short-range process. Two novel effects that separated MAE from motion detection are described, and suggestions for exploring them are outlined.     

Blocking of spatial learning between enclosure geometry and a local landmark

In a virtual environment, blocking of spatial learning to locate an invisible target was found reciprocally between a distinctively shaped enclosure and a local landmark within its walls. The blocking effect was significantly stronger when the shape of the enclosure rather than the landmark served as the blocking cue. However, the extent to which the landmark blocked enclosure-shape learning was not influenced by increasing the physical salience of the landmark. The outcomes are the first to suggest that cue-interaction effects, commonly found in human and animal contingency learning experiments, are also found in human spatial learning based on landmarks and enclosure walls. The data are discussed in terms of spatial reference frames.     

Interoceptive awareness and unaware fear conditioning: are subliminal conditioning effects influenced by the manipulation of visceral self-perception?

Research has shown repeatedly that attention influences implicit learning effects. In a similar vein, interoceptive awareness might be involved in unaware fear conditioning: The fact that the CS is repeatedly presented in the context of aversive bodily experiences might facilitate the development of conditioned responding. We investigated the role of interoceptive attention in a subliminal conditioning paradigm. Conditioning was embedded in a spatial cueing task with subliminally presented cues that were followed by a masking stimulus. Response times to the targets that were either validly or invalidly predicted by the cues served as index of conditioning. Interoceptive attention was manipulated between-subjects. Half the participants completed a heartbeat detection task before conditioning. This task tunes attention to one’s own bodily signals. We found that conditioned responding was facilitated in this latter group of participants. These results are in line with the hypothesis that a rise interoceptive attention enhances unaware conditioned responding.     

No evidence for overshadowing or facilitation of spatial pattern learning by visual cues

Two experiments were conducted to examine the effects of redundant and relevant visual cues on spatial pattern learning. Rats searched for hidden food items on the tops of poles that formed a square (Experiment 1) or a checkerboard (Experiment 2) pattern. The experimental groups were trained with visual cues that specified the locations of the baited poles. All groups were tested without visual cues so that any overshadowing or facilitation of spatial pattern learning by visual cues could be detected. Spatial choices were controlled by the spatial pattern and by the visual cues in both experiments. However, there was no evidence of overshadowing or facilitation of spatial pattern learning by visual cues in either experiment. The results are consistent with the idea that the representation of the spatial pattern that guides choices is not controlled by the same learning processes as those that produce associations between visual cues and food locations.     

Interference with ingestional aversion learning produced by preexposure to the unconditioned stimulus: Associative and nonassociative aspects

Preconditioning exposure to a lithium chloride unconditioned stimulus (US) interferes with the subsequent conditioning of a taste aversion. Multiple US preexposures produce a long-lasting or durable interference with aversion learning, whereas a single US exposure produces a transient interference effect. The present experiments demonstrate that the durable US preexposure effect is substantially reduced if the method of drug treatment (infusion versus injection of the drug into the peritoneal cavity) or the spatial cues present during drug treatment (home cage versus a distinctive environment) are changed between the preexposure and taste conditioning phases of the experiment. In contrast, these manipulations do not attenuate the proximal/transient US preexposure effect. These findings indicate that different mechanisms are responsible for the two US preexposure effects. The results are consistent with previous suggestions that the durable effects of lithium preexposure are due to associative interference produced by the exteroceptive stimuli that accompany drug administrations and indicate that the proximal/transient US preexposure effect is mediated by nonassociative mechanisms.     

Effects of US-Alone Presentations on Human Evaluative Conditioning

In the context of evaluative conditioning, the effects of additional presentations of the unconditioned stimulus (US) prior to conditioning (US preexposure) or after conditioning (US postexposure) were examined using between- and within-subjects control conditions. Two experiments that differed with respect to the nationality of the subjects were conducted. In both experiments, US-alone presentations reduced the magnitude of the evaluative response. The US pre- and postexposure effects were observed in subjects classified as aware as well as in subjects classified as unaware of the experimental contingencies. Another finding is that the evaluative conditioning procedure described by Martin and Levey (1978; Levey & Martin, 1975) resulted in reliable conditioning effects also in an American sample, thus extending the scope of that special evaluative conditioning paradigm. The findings are discussed in the context of recent models of classical and evaluative conditioning. Especially, the unexpected US postexposure effect gives rise to speculations concerning the learning process underlying evaluative conditioning.     

Roles of egocentric and allocentric spatial representations in locomotion and reorientation

Four experiments investigated the nature of spatial representations used in locomotion. Participants learned the layout of several objects and then pointed to the objects while blindfolded in 3 conditions: before turning (baseline), after turning to a new heading (updating), and after disorientation (disorientation). The internal consistency of pointing in the disorientation condition was relatively high and equivalent to that in the baseline and updating conditions, when the layout had salient intrinsic axes and the participants learned the locations of the objects on the periphery of the layout. The internal consistency of pointing was disrupted by disorientation when participants learned the locations of objects while standing amid them and the layout did not have salient intrinsic axes. It was also observed that many participants retrieved spatial relations after disorientation from the original learning heading. These results indicate that people form an allocentric representation of object-to-object spatial relations when they learn the layout of a novel environment and use that representation to locate objects around them. Egocentric representations may be used to locate objects when allocentric representations are not of high fidelity.