Chronic stress modulates the use of spatial and stimulus-response learning strategies in mice and man

Acute stress modulates multiple memory systems in favor of caudate nucleus-dependent stimulus-response and at the expense of hippocampus-dependent spatial learning and memory. We examined in mice and humans whether chronic stress has similar consequences. Male C57BL/6J mice that had been repeatedly exposed to rats ("rat stress") used in a circular hole board task significantly more often a stimulus-response strategy (33%) than control mice (0%). While velocity was increased, differences in latency to exit hole, distance moved or number of holes visited were not observed. Increased velocity and performance during retention trials one day later indicates altered emotionality and motivation to explore in rat stressed mice. Forty healthy young men and women were split into "high chronic stress" and "low chronic stress" groups based on their answers in a chronic stress questionnaire ("Trier Inventory of Chronic Stress"-TICS) and trained in a 2D task. A test trial immediately after training revealed that participants of the "high chronic stress" group used the S-R strategy significantly more often (94%) than participants of the "low chronic stress" group (52%). Verbal self-reports confirmed the strategy derived from participants' choice in the test trial. Learning performance was unaffected by the chronic stress level. We conclude that one consequence of chronic stress is the shift to more rigid stimulus-response learning, that is accompanied by changes in motivational factors in mice.     

Sequence learning in Parkinson's disease: the effect of spatial stimulus-response compatibility

Patients with Parkinson's disease (PD) have repeatedly demonstrated reduced sequence-specific learning effects in serial reaction time tasks (SRTs). Previous research with PD patients has mainly employed the 'classical' SRT task, involving a spatially compatible assignment of stimuli and responses. From cognitive research, it is known that spatial compatibility triggers rapid, automatic responses in the direction of the stimulus. Automatic responding has shown to be disinhibited in PD patients and may therefore interfere with stimulus anticipation during the learning process. The aim of the present study was to test this hypothesis by investigating if reduced sequence-specific learning depends on spatial stimulus-response compatibility. PD patients and age-matched controls were examined either with an SRT variant involving central stimulus presentation, thereby preventing automatic linking of stimulus and response locations, or with a spatially compatible SRT task. Patients showed reduced sequence-specific learning effects only when the stimulus-response assignment was spatially compatible. This pattern of results confirms the hypothesis that sequence learning deficits in PD may result from a predominance of automatic response activation over learning-based stimulus anticipations during the learning phase.     

Conditional probability versus spatial contiguity in causal learning: Preschoolers use new contingency evidence to overcome prior spatial assumptions

This study examines preschoolers' causal assumptions about spatial contiguity and how these assumptions interact with new evidence in the form of conditional probabilities. Preschoolers saw a toy that activated in the presence of certain objects. Children were shown evidence for the toy's activation rule in the form of patterns of probability: The toy was more likely to activate either when objects made contact with its surface (on condition) or when objects were several inches above its surface (over condition). In Experiment 1, 61 three-year-olds saw a deterministic activation rule. In Experiments 2 and 3, 48 four-year-olds saw an activation rule that was probabilistic. In Experiment 4, 30 four-year-olds saw a screening-off pattern of activation. In all 4 experiments, children used new evidence in the form of patterns of probability to make accurate causal inferences, even in the face of conflicting prior beliefs about spatial contiguity. However, children were more likely to make correct inferences when causes were spatially contiguous, particularly when faced with ambiguous evidence.     

Functional networks involved in spatial learning strategies in middle-aged rats

Our aim was to assess the way that middle-aged rats solve spatial learning tasks that can be performed using different strategies. We assessed the brain networks involved in these spatial learning processes using Principal Component Analysis. Two tasks were performed in a complex context, a four-arm radial maze, in which each group must use either an allocentric or an egocentric strategy. Another task was performed in a simple T-maze in which rats must use an egocentric strategy. Brain metabolic activity was quantified to evaluate neural changes related to spatial learning in the described tasks. Our findings revealed that two functional networks are involved in spatial learning in aged rats. One of the networks, spatial processing, is composed of brain regions involved in the integration of sensory and motivational information. The other network, context-dependent processing, mainly involves the dorsal hippocampus and is related to the processing of contextual information from the environment. Both networks work together to solve spatial tasks in a complex spatial environment.     

Priming and stimulus-response learning in perceptual classification tasks

Participants often respond more quickly and more accurately to a repeated stimulus compared to a non-repeated one, a phenomenon known as repetition priming. In semantic classification tasks priming appears to be largely attributable to the learning of stimulus-decision and stimulus-response associations, which allow participants to bypass many of the processes engaged during initial stimulus analysis. The current study tested whether stimulus-response learning plays a similarly dominant role in priming that occurs in perceptual classification tasks. Unfamiliar objects were used as stimuli to reduce the influence of semantic processes on priming and the task switched for all test trials to eliminate stimulus-decision learning. The results showed across-task priming as measured by reaction time facilitation and improved accuracy when the response remained the same during the encoding and test phases. When the response switched, similar levels of reaction time facilitation were observed, but priming as measured by accuracy was significantly reduced and no longer significant. These findings indicate that stimulus-response learning contributes to priming in perceptual classification tasks, but does not play a dominant role. Significant stimulus-level learning that is independent of the task and response also occurs and likely indexes facilitated perceptual processing of the objects.     

Multiple spatial learning strategies in goldfish (Carassius auratus)

There is a considerable amount of evidence that mammals and birds can use different spatial learning strategies based on multiple learning and memory systems. Unfortunately, only a few studies have investigated spatial learning and memory mechanisms in other vertebrates. This study aimed to identify the strategies used by goldfish to solve two different spatial tasks in a series of three experiments. In experiment 1, two groups of goldfish (Carassius auratus) were trained either in a spatial constancy task (SC), in which visual cues signalled the goal indirectly, or in a directly cued task (DC) in which similar cues signalled the goal directly. Transfer tests were conducted to study the effects of discrete cue deletion on the performance in both tasks. In these transfer tests the performance of the animals trained in the DC task dropped to chance level when the cue that signalled the goal directly was removed. In contrast, the removal of any single cue did not disrupt SC performance. In experiment 2, fish trained in the SC or the DC task were trained with the goal reversed. Goldfish in the SC group needed fewer sessions to master the reversal task than DC animals. Finally, experiment 3 investigated the effects of a substantial modification of the geometrical features of the apparatus on the performance of animals trained in the SC or in the DC condition. The performance of DC goldfish was not affected, whereas the same change disrupted performance in the SC animals despite the presence of the visual cues. These results suggest that there are separate spatial learning and memory systems in fish. Whereas the DC animals used a typical guidance strategy, relying only on the cue that signalled the goal directly, SC fish relied on a strategy with the properties of an actual spatial mapping system. Thus, the comparative approach points to the generality of these learning strategies among vertebrates. Received: 10 October 1998 / Accepted after revision: 16 April 1999     

Interaction between locale and taxon strategies in human spatial learning

Three computer-based experiments which tested human participants in a non-immersive virtual watermaze task sought to determine factors which dictate whether the presence of a visual platform disrupts locale learning and taxon learning. In Experiment 1, the visible platform disrupted locale but not taxon learning based on viewpoint-independent and viewpoint-dependent information, respectively. In Experiment 2, taxon learning based on non-geometric cues providing viewpoint-dependent information was disrupted by the visible platform when the cues required relational information to disambiguate them from other cues. Experiment 3 placed participants in an isosceles triangular pool. The presence of the visible platform did not disrupt the encoding of relational information provided by shape of the pool. These results support the notion that geometric cues are encoded in a separate module which is impenetrable to non-geometric cues not creating the shape of the environment.     

Imitative learning of stimulus-response and response-outcome associations in pigeons

A novel automated procedure was used to study imitative learning in pigeons. In Experiments 1 and 2, observer pigeons witnessed a demonstrator pigeon successfully performing an instrumental discrimination in which different discriminative stimuli indicated which of 2 topographically distinct responses (R1 and R2) resulted in the delivery of seed. The observers were then presented with the discriminative stimuli and given access to the response panel. Observer pigeons' behavior during the discriminative stimuli was influenced by how the demonstrator had responded during these stimuli. In Experiment 3, observers witnessed demonstrator pigeons performing R1 for Outcome 1 and R2 for Outcome 2. Observers then received a procedure designed to devalue Outcome 1 relative to Outcome 2 and were subsequently less likely to perform R1 than R2. These results suggest that pigeons can learn both stimulus response and response-outcome associations by observation.     

Effects of multiple reference points in spatial stimulus-response compatibility.

In two experiments, spatial stimulus-response compatibility effects in situations where the stimulus could appear in eight different locations were investigated. The locations were obtained as a result of orthogonal manipulation of hemispace, visual hemifield within hemispace, and relative position within hemifield. In the first experiment, only relative position within hemifield was relevant for selecting one of two responses (left or right). The results showed that both hemifield and relative position formed the basis of compatibility effects. In the second experiment, which was in most respects identical to the first, all spatial information was irrelevant. Only the geometrical shape of the stimulus determined the correct response. The results showed three S-R compatibility effects, based on hemispace, hemifield, and relative position. These results contradict earlier findings, and have implications for models of stimulus-response compatibility.     

The use of computer-controlled video disks in the study of spatial learning

The empirical study of spatial learning has been constrained by a range of methodological and technical limitations. Part of these are due to constraints in the stimulus delivery systems for pictorial displays that are available to the researcher. Surrogate travel systems—based on model photography, off-line computer graphics, and on-line computer graphics—provide alternative research settings. Yet none of the systems provides for both detailed displays and ease of subject interaction, both of which prove important in the investigation of spatial learning. Computer-controlled video-disk systems do allow both detail and interaction, therefore providing a research tool in which spatial learning phenomena can be observed carefully. A pilot study of spatial learning using a video-disk system is briefly described to demonstrate the sorts of phenomena observable with such a system.     

Cross language learning disabilities and verbal versus spatial memory

The research reported in this talk involves comparisons of verbal and spatial memory tasks across groups of children (and adults) with different types of learning difficulties. The research focuses on children with literacy acquisition problems and investigates whether such problems are related to specific areas of deficit. In the first piece of research, children with dyslexia (literacy learning problems) and dyspraxia (motor deficits) were contrasted on measures of memory (for example, tasks that required the retention of sequences of verbal material or spatial movements) and additional measures of literacy (reading and spelling), phonological (awareness of sounds within words) and motor (fine and gross motor tasks) functioning. The data were consistent with a dissociation between tasks/groups such that dyslexics showed weak phonological processing but intact visuo-spatial processing, whereas children with dyspraxia showed weaknesses on task involving visuo-spatial information, but average levels of performance on tasks that required phonological processing. Similar results were identified amongst adult groups, consistent with a deviant level of functioning rather than a developmental delay. A second line of research contrasted children with or without literacy problems across language backgrounds (English, Arabic, Chinese and bilingual children). Consistent with the dyslexia data, children with poor English literacy skills showed weaknesses in verbal/phonological memory tasks but not in visuo-spatial memory. However, for Chinese-language children, visuo-spatial memory differed between good and poor literacy learners, but there was little evidence for verbal memory differences. In contrast, the Arabic and bilingual children showed differences in both verbal and visuo-spatial areas, although the evidence was consistent with enhanced visual/spatial skills amongst the good literacy groups, rather than poor literacy children showing weaknesses in those tasks. These data suggest that the influence of memory skills on learning may vary with the language of instruction. A final line of enquiry considers whether teaching strategies to children with learning difficulties may overcome some of the identified memory deficits and lead to better levels of learning. English language children with learning difficulties were taught visual and verbal strategies to support retention of materials in short-term memory tasks. In the majority of cases, learning was improved when it focused on visuo-spatial strategies but not when verbal strategies were used. These data support the relationship between learning difficulties and different aspects of short-term memory that may lead to poor levels of learning. It also presents evidence that memory (particularly those related to visuo-spatial) processes are influenced by the context within which learning is taking place, both in terms of the language of instruction and the strategies used to support learning. For some children with educational difficulties based around language-related deficits, visuo-spatial strategies may support acquisition.     

Unintentional and intentional learning of noncorresponding stimulus-response associations in the Simon task

The Simon effect is a robust phenomenon that persists after extensive practice. However, several studies using a transfer paradigm have shown that the Simon effect is eliminated after practicing a location-relevant task with an incompatible spatial mapping. The present study examined whether this transfer effect is a result of implicit, procedural knowledge developed through repeated execution of noncorresponding responses in the practice session or a consequence of explicitly learning and reinstating a noncorresponding mapping rule. Results from two experiments show that, although a small part of the transfer effect may be due to residual activation of noncorresponding S-R associations from the prior task, the larger and more stable part is likely due to response-selection strategies performed intentionally in the practice task.     

A single bout of aerobic exercise modulates motor learning performance and cortical excitability in humans

BackgroundEvidence indicates beneficial effects of aerobic exercise on motor learning performance, which might be caused by the impact of aerobic exercise on cortical excitability. It is thus suggested that physiological effects of aerobic exercise on cortical excitability determine the effects of aerobic exercise on motor learning. Nevertheless, respective results usually come from independent studies, and a prove of the causal relationship between neurophysiological and motor learning effects is still missing. This study aims to explore the impact of a single bout of aerobic exercise on brain physiology and motor learning, and the association between these phenomena in humansMethodThe study was conducted in a cross-over design. In twenty healthy subjects, cortical excitability and motor learning were assessed before and after a single bout of aerobic exercise or a control interventionResultsThe results show that aerobic exercise improved motor sequence learning and enhanced cortical excitability in humans. Furthermore, a correlation between the exercise-dependent alteration of cortical excitability (short intracortical inhibition, which is determined primarily by the GABAergic system) and improvement of motor learning has been foundDiscussionThe study found motor learning performance-improving effects of aerobic exercise, and these results might be explained by an exercised-caused alteration of cortical excitability, especially a reduction of GABA activity.     

Function learning: induction of continuous stimulus-response relations.

This research investigates how people learn to select particular response magnitudes along one physical dimension (duration) when given stimulus magnitudes from another dimension (spatial extent). Stimuli and correct responses were related by either a power function, a logarithmic function, or a linear function with a positive intercept. The power function was learned quickly and accurately. In contrast, systematic response biases occurred during the early phases of learning the logarithmic and linear functions. As practice progressed, however, the biases gradually disappeared. These results support an adaptive regression model. According to it, people learn functions through a subjective process analogous to statistical regression. There is assumed to be an initial constraint that treats stimulus-response pairs as if a power function characterizes them.     

A psychophysiological study of the use of partial information in stimulus-response translation.

Ss performed a hybrid go/no-go reaction task in which colored letters were assigned in various ways to 4 finger responses, 2 on each hand. In addition to reaction time, psychophysiological measures were used to assess the duration of stimulus identification and the onset of central and peripheral motor activity. The results suggest that response selection can begin on the basis of 1 stimulus dimension (e.g., color), while the other dimension (e.g., letter form) has not yet been identified. Other results are discussed with regard to "selection for action" (Allport, 1987) and the importance of stimulus-response translation strategies in the use of partial information.     

Four-dimensional spatial reasoning in humans

Human subjects practiced navigation in a virtual, computer-generated maze that contained 4 spatial dimensions rather than the usual 3. The subjects were able to learn the spatial geometry of the 4-dimensional maze as measured by their ability to perform path integration, a standard test of spatial ability. They were able to travel down a winding corridor to its end and then point back accurately toward the occluded origin. One interpretation is that the brain substrate for spatial navigation is not a built-in map of the 3-dimensional world. Instead it may be better described as a set of general rules for manipulating spatial information that can be applied with practice to a diversity of spatial frameworks.     

Anticipatory response control in motor sequence learning: evidence from stimulus-response compatibility

Three experiments using a serial four-choice reaction-time (RT) task explored the interaction of sequence learning and stimulus-based response conflict. In Experiment 1, the spatial stimulus-response (S-R) mapping was manipulated between participants. Incompatible S-R mappings produced much higher RTs than the compatible mapping, but sequence learning decreased this S-R compatibility effect. In Experiment 2, the spatial stimulus feature was made task-irrelevant by assigning responses to symbols that were presented at unpredictable locations. The data indicated a Simon effect (i.e., increased RT when irrelevant stimulus location is spatially incompatible with response location) that was reduced by sequence learning. However, this effect was observed only in participants who developed an explicit sequence representation. Experiment 3 replicated this learning-based modulation of the Simon effect using explicit sequence-learning instructions. Taken together, the data support the notion that explicit sequence learning can lead to motor 'chunking', so that pre-planned response sequences are shielded from conflicting stimulus information.     

Attention modulates the learning of multiple contingencies

We employed a novel version of the serial reaction time task to test the idea that human implicit learning allows the simultaneous learning of multiple independent contingencies and that this learning may proceed in the absence of attention. Using probabilistic sequences, we showed that both a primary sequence (the focus of the experimental task) and a statistically independent secondary sequence could be learned across 4,800 target localization trials, provided the perceptual load of the primary task was low. However, learning of the secondary sequence was abolished under conditions of high perceptual load. These findings suggest that there are attentional limitations on the learning of multiple contingencies.