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Spatial and nonspatial escape strategies in the Barnes maze.

The Barnes maze is a spatial memory task that requires subjects to learn the position of a hole that can be used to escape the brightly lit, open surface of the maze. Two experiments assessed the relative importance of spatial (extra-maze) versus proximal visible cues in solving the maze. In Experiment 1, four groups of mice were trained either with or without a discrete visible cue marking the location of the escape hole, which was either in a fixed or variable location across trials. In Experiment 2, all mice were trained with the discrete visible cue marking the target hole location. Two groups were identical to the cued-target groups from Experiment 1, with either fixed or variable escape locations. For these mice, the discrete cue either was the sole predictor of the target location or was perfectly confounded with the spatial extra-maze cues. The third group also used a cued variable target, but a curtain was drawn around the maze to prevent the use of spatial cues to guide navigation. Probe trials with all escape holes blocked were conducted to dissociate the use of spatial and discrete proximal cues. We conclude that the Barnes maze can be solved efficiently using spatial, visual cue, or serial-search strategies. However, mice showed a strong preference for using the distal room cues, even when a discrete visible cue clearly marked the escape location. Importantly, these data show that the cued-target control version of the Barnes maze as typically conducted does not dissociate spatial from nonspatial abilities.

Analysis of Variance↗

Exposure to stable flies reduces spatial learning in mice: involvement of endogenous opioid systems.

Biting flies influence both the physiology and behaviour of domestic and wild animals. This study demonstrates that relatively brief (60 min) exposure to stable flies, Stomoxys calcitrans (L.), affects the spatial abilities of male mice. Stable fly exposure resulted in poorer subsequent performance in a water maze task in which individual mice had to learn the spatial location of a submerged hidden platform using extramaze visual cues. Determinations of spatial acquisition and retention were made with mice that had been previously exposed for 60 min to either stable flies or house flies, Musca domestica (L.). Mice exposed to stable flies displayed over one day of testing (six blocks or sets of four trials) significantly poorer acquisition and retention of the water maze task than either mice that had been exposed to house flies or fly-naive mice. This attenuation of spatial learning occurred in the absence of any evident sensorimotor or motivational impairments. The reduction in spatial abilities involved endogenous opioid systems, as the decreased performance resulting from stable fly exposure was blocked by pre-treatment with the prototypic opiate antagonist, naltrexone. These results indicate that relatively brief exposure to biting flies can lead to a decrease in spatial abilities which is associated with enhanced endogenous opioid activity. These results support the involvement of endogenous opioid systems in the mediation of the behavioural and physiological effects of biting fly exposure. They further suggest that decreases in spatial abilities and performance may be part of the behavioural consequences of biting fly exposure in domestic and wild animals.

Animals↗

Activity of hippocampal formation neurons in the monkey related to a conditional spatial response task.

To analyze neurophysiologically the functions of the primate hippocampus, the activity of 905 single hippocampal formation neurons was analyzed in two rhesus monkeys performing a conditional spatial response task known to be impaired in monkeys and in man by damage to the hippocampus or fornix. In the task, the monkey learned to make one spatial response, touching a screen three times when he saw one visual stimulus on the video monitor, and a different spatial response, of withdrawing his hand from the screen, when a different visual stimulus was shown. Fourteen percent of the neurons fired differentially to one or the other of the stimulus-spatial response associations. The mean latency of these differential responses was 154 +/- 44 (SD) ms. The firing of these neurons was shown to reflect a combination of the particular stimulus and the particular response associated by learning in the stimulus-response association task and could not be accounted for by the motor requirements of the task, nor wholly the stimulus aspects of the task, as demonstrated by testing their firing in related visual discrimination tasks. Responsive neurons were found throughout the hippocampal formation, but were particularly concentrated in the subicular complex and the CA3 subfield. These results show that single hippocampal neurons respond to combinations of the visual stimuli and the spatial responses with which they must become associated in conditional spatial response tasks and are consistent with the suggestion that part of the mechanism of this learning involves associations between visual stimuli and spatial responses learned by single hippocampal neurons.

Action Potentials↗

Brain-behavior relationships: evidence from practice effects in spatial stimulus-response compatibility.

1. We measured relative cerebral blood flow (rCBF) changes with positron emission tomography and H(2) 15O in six normal subjects repeatedly performing a spatial stimulus-response compatibility task. Subjects had two motor response conditions. They were instructed to respond with the left hand to a left visual field light stimulus and with the right hand to a right visual field light stimulus (compatible condition), and with the right hand to a left visual field light stimulus and with the left hand to a right visual field light stimulus (incompatible condition). Six rCBF measurements per condition were performed in each subject. 2. Reaction times were faster (P < 0.0005) in the compatible (287 ms) than the incompatible (339 ms) condition (spatial stimulus-response compatibility effect). A bilateral increase (P < 0.05) in rCBF in the superior parietal lobule of the two hemispheres was observed during the incompatible condition when compared with the compatible one. No rCBF decreases were observed. Reaction times correlated (P < 0.0001) with the rCBF in the two activated superior parietal lobule areas. 3. Reaction times decreased with practice according to a linear trend (P < 0.05). Practice-related linear rCBF increases (P < 0.05) were observed in the dorsolateral prefrontal, premotor, and primary motor cortex of the left hemisphere. No significant rCBF decreases were observed. 4. Practice did not affect the spatial stimulus-response compatibility effect. A parallel shortening of reaction times was observed in both compatible and incompatible conditions, in both left and right hand responses, and in both left and right visual fields. Accordingly, when rCBF was analyzed, the spatial stimulus-response compatibility by practice interaction did not show any significant activated area. 5. These findings suggest that the two activated areas in the left and right superior parietal lobules subserve the mapping of the visual stimulus spatial attributes onto the motor response spatial attributes and that the rCBF increases in the incompatible response condition represent the more complex computational remapping required when stimuli and response do not match spatially. 6. The dorsolateral prefrontal, premotor, and motor rCBF linear increases in the left hemisphere seem to reflect the effect of practice on cortical processes common to both compatible and incompatible response conditions. These cortical processes presumably strengthen the links between stimuli and responses under different stimulus-response compatibility conditions. The lateralization of the rCBF increases suggests a left hemisphere superiority in these processes.

Adult↗

Relational spatial reasoning by a nonhuman: the example of capuchin monkeys.

The authors review spontaneous manipulation and spatial problem solving by capuchin monkeys to illuminate the nature of relational reasoning (wherein two or more elements of a problem or situation are considered together to arrive at a course of action) that these monkeys use in goal-directed activity. Capuchin monkeys master problems with one, two, or three spatial relations, and if more than one relation, at least two relations may be managed concurrently. They can master static and dynamic relations and, with sufficient practice, can produce specific spatial relations through both direct and distal action. Examining capuchins' spatial problem-solving behavior with objects in the framework of a spatial relational reasoning model leads to new interpretations of previous studies with these monkeys and other nonhuman animals. The model produces a variety of testable predictions concerning the contribution of relational properties to spatial reasoning. It also provides conceptual linkages with neurological processes and cognitive analyses of physical reasoning. Understanding relational spatial reasoning, including tool use, in a wider view is vital to informed, principled comparison of problem solving and the use of technology across species, across ages within species, and across eras in human prehistory.

Animals↗

Spatial articulation affects lightness.

In a recent paper, Agostini and Bruno (1996) showed that the size of simultaneous lightness contrast increases under Gelb lighting. To extend Agostini and Bruno's work, we applied their methodology to a set of more spatially articulated displays. In four experiments, we investigated the role of spatial articulation on the size of the simultaneous lightness contrast effect. In the first experiment, we found a decrease of the simultaneous lightness contrast effect as the spatial articulation increased. In the second experiment (the control experiment), performed under homogeneous illumination, we found that the effect of spatial articulation is not detectable, even though the data seem to show the same trend as that in the previous experiment. In the third experiment, we found that spatial articulation affects not only the middle reflectance region, but also the lowest one. As the spatial articulation increases, the effect on the lightnesses of both regions decreases. In the last experiment, performed with a reduced range of reflectances, we found a lightening effect for all the reflectances and, again, an effect of spatial articulation. The results of these experiments are interpreted according to the model proposed by Gilchrist et al. (in press).

Humans↗

Involvement of brain-derived neurotrophic factor in spatial memory formation and maintenance in a radial arm maze test in rats.

Brain-derived neurotrophic factor (BDNF) regulates both short-term synaptic functions and activity-dependent synaptic plasticity such as long-term potentiation. In the present study, we investigated the role of BDNF in the spatial reference and working memory in a radial arm maze test. The radial arm maze training resulted in a significant increase in the BDNF mRNA expression in the hippocampus, although the expression in the frontal cortex did not change. When spatial learning was inhibited by treatment with 7-nitroindazole, an inhibitor of brain nitric oxide synthase, the increase in the hippocampal BDNF mRNA did not occur. To clarify the causal relation between BDNF mRNA expression and spatial memory formation, we examined the effects of antisense BDNF treatment on spatial learning and memory. A continuous intracerebroventricular infusion of antisense BDNF oligonucleotide resulted in an impairment of spatial learning, although the sense oligonucleotide had no effect. Treatment with antisense, but not sense, BDNF oligonucleotide was associated with a significant reduction of BDNF mRNA and protein levels in the hippocampus. Furthermore, treatment with antisense BDNF oligonucleotide in rats, which had previously acquired spatial memory by an extensive training, impaired both reference and working memory. There were no differences in locomotor activity, food consumption, and body weight between the antisense and sense oligonucleotide-treated rats. These results suggest that BDNF plays an important role not only in the formation, but also in the retention and/or recall, of spatial memory.

Analysis of Variance↗

Long-term enhancement of hippocampal synaptic transmission and the acquisition of spatial information.

The hypothesis that memories are stored as a specific distribution of strengths in a population of modifiable synapses was examined by the bilateral induction of long-term enhancement in synapses of the main afferent fiber system to the hippocampal formation in rats. Brief, high-frequency activation of the perforant pathway in chronically prepared animals resulted in a persistent increase in the field EPSP and population spike, measured extracellularly in fascia dentata. This treatment resulted in a profound and persistent deficit in the acquisition of new spatial information in a task requiring spatial "reference" memory, and disruption of recently acquired spatial information. Well-established spatial memory was completely unaffected, however, as was the acquisition of spatial information into short-term "working" memory. These results support the hypothesis that, during the formation of "cognitive maps," spatial information must be temporarily stored at modifiable synapses at the input stage to the hippocampal formation, but that this information is not needed once the representation of the environment is well established. Spatial working memory, in a familiar environment, appears not to depend on the distribution of synaptic strengths in this system at all.

Animals↗

Spatial and temporal processing of threshold data for detection of progressive glaucomatous visual field loss.

OBJECTIVE: To evaluate the effect of spatial and temporal filtering of threshold visual field data on the ability of pointwise linear regression (PLR) to detect progressive glaucomatous visual field loss. METHODS: Longitudinal visual field data (Full-Threshold Program 30-2 test point pattern) were simulated using a computer model of glaucomatous visual field progression. This approach permitted construction of a "gold standard" because matching visual field data without variability could be generated and analyzed. Four clustered progressive defects were produced, consisting of 2, 3, 9, and 18 locations, respectively, each with progression rates of -1 and -2.5 dB/y. Pointwise linear regression was used to identify progressive test locations (criterion for progression of statistically significant slope of < or =-1 dB/y, P<.05). Each visual field series was analyzed after the following 3 procedures: (1) no filtering (unprocessed data), (2) Gaussian spatial possessing (3 x 3 grid), and (3) temporal processing (2 field moving average). The effect of spatial and temporal processing on PLR discriminatory power for progression detection was quantified by comparison with the gold standard. RESULTS: Spatial processing reduced PLR sensitivity to levels below that achieved for analysis of unprocessed data for small progressive defects (< or =9 locations) or at the low true progression rate (-1 dB/y). Under these conditions, spatial processing caused small PLR specificity improvement. Spatial processing only improved PLR sensitivity above unprocessed levels when progressive defects were large and changing rapidly (progression rate of -2.5 dB/y). Temporal processing gave consistent PLR improvement in sensitivity for all defect sizes and true progression rates. Pointwise linear regression sensitivity gain provided by temporal processing allowed progression to be detected 2 to 3 visual fields earlier than for analysis of raw data. Specificity dropped slightly as a result of temporal processing but remained at 89% or above for all conditions studied. CONCLUSIONS: Gaussian spatial processing reduces PLR discriminatory power with low true progression rates or small progressive defect sizes and, therefore, is of limited use for detection of progressive visual field loss. Temporal processing improves the sensitivity of PLR and reduces the number of tests required to detect progressive loss with minimal loss of specificity. CLINICAL RELEVANCE: Image processing techniques can be applied to threshold visual field data to enhance sensitivity or specificity of PLR for the determination of progressive change. This investigation demonstrates that temporal processing may assist with the detection of significant progressive visual field loss with fewer test results than unprocessed data.

Computer Simulation↗

Spatial patterns of human gene frequencies in Europe.

The aims of this study of spatial patterns of human gene frequencies in Europe are twofold. One is to present new methodology developed for the analysis of such data. The other is to report on the diversity of spatial patterns observed in Europe and their interpretation as evidence of population processes. Spatial variation in 59 allele and haplotype frequencies (26 genetic systems) for polymorphisms in blood antigens, enzymes, and proteins is analyzed for an aggregate of 3,384 localities, using homogeneity tests, one-dimensional and directional spatial correlograms, and SYMAP interpolated surfaces. The data matrices are reduced to reveal the principal patterns by clustering techniques. The findings of this study can be summarized as follows: 1) There is significant heterogeneity in allele frequencies among the localities for all but one genetic system. 2) There are significant spatial patterns for most allele frequencies. 3) There is a substantial minority of clinal patterns in these populations. Clinal trends are found more frequently in HLA alleles than for other variables. North-south and northwest-southwest gradients predominate. 4) There is a strong decline in overall genetic similarity with geographic distance for most variables. 5) There are few, if any, appreciable correlations in pairs of allele frequencies over the continent, and there is little interesting correlation structure in the resulting correlation matrix. 6) Few spatial correlograms are markedly similar to each other, yet they form well-defined clusters. Spatial variation patterns, therefore, differ among allele frequencies. Patterns of human gene frequencies in modern Europe are diverse and complex. No single model suffices for interpretation of the observed genetic structure. Some clinal patterns reported here support the Neolithic demic-expansion hypothesis, others suggest latitudinal selection. Most of the clinal patterns are in HLA alleles, but there is also evidence from ABO for east-west migration diffusion. The majority of patterns are patchy, consistent with hypotheses of isolation by distance or of settlement of genetically differing, subsequently expanding ethnic groups. While undoubtedly there has been an ongoing stochastic process of differentiation consistent with the isolation-by-distance model, this has not obscured the directional patterns caused by migration (demic diffusion), and has perhaps only reinforced the contribution from settlement of ethnic units to patterns of genetic variation. However, the impact of the latter is most difficult to discern and requires further methodological developments.

ABO Blood-Group System↗

Hierarchical Bayesian spatial modelling of small-area rates of non-rare disease.

We present Bayesian hierarchical spatial models for the analysis of the geographical distribution of a non-rare disease or event. The work is motivated by the need for ascertaining regional variations in health services outcomes and resource use and for assessing the potential sources of these variations. The models discussed herein readily accommodate random spatial effects and covariate effects. We discuss Bayesian inferential framework and implementation of a hybrid Markov chain Monte Carlo method for full Bayesian model inference. The methods are illustrated through an analysis of regional variation in chronic lung disease (CLD) rates among neonatal intensive care unit (NICU) patients across Canada. Specifically, we first present a random effects binomial model for spatially correlated CLD rates, with random spatial effects accounting for latent or covariate effects. These random spatial effects depict regional or spatial variation in chronic lung disease occurrence. We then extend this model to include covariates. With this extension, we assess residual spatial effects and the extent to which risk factors such as illness severity at NICU admission, low birth weight, and very low birth weight influence the CLD rate variation.

Bayes Theorem↗

Spatial and reversal learning in congeneric lizards with different foraging strategies.

Environmental demands that require intensive search for mates, food and nest sites are correlated with efficient spatial memory in many mammalian and avian species. This convergence of evidence has led to the view that spatial memory, and the neurological structures associated with it, have been selected in niches that require memory for the location of goal objects. Whether such evolutionary demands are also correlated with nonspatial abilities that require flexible use of associations similar to those required for spatial memory has not been well studied. In addition, correlations between niche types and the use of spatial or nonspatial memory have not been investigated in nonmammalian, nonavian taxa. In this study, we investigated the relationship between foraging strategies and performance on two tasks, one spatial and the other nonspatial, in congeneric lizard species: Acanthodactylus boskianus, an active forager that collects clumped sedentary prey, Acanthodactylus scutellatus, a sit-and-wait predator that collects distributed mobile prey. The two species did not differ in their performance of a spatial memory task, but A. boskianus, the active forager, performed better on the reversal of a visual discrimination, a nonspatial task. These findings question the generality of the spatial adaptation model for vertebrates. We present the pliancy hypothesis, which we developed to account for these results. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

On spatial frequencies and cerebral hemispheres: some remarks from the electrophysiological and neuropsychological points of view.

The spatial frequency hypothesis on hemispheric specialization gave rise to contradictory experimental results, commented on in Brain and Cognition by Christman (1989) and Peterzell (1991). The question is discussed through a review of the electrophysiological and neuropsychological research on hemispheric asymmetry of spatial frequency processing. The general hypothesis of the hemispheric specialization for this basic visual information appears to be supported by recent works on evoked potentials by gratings and checkerboards. However, an interaction between the cerebral hemisphere, spatial frequency, and temporal frequency was found more than a sharp dichotomy between low (right hemisphere) and high spatial frequencies (left hemisphere), as indeed it was proposed by the spatial frequency hypothesis. Other relevant physical parameters in generating the hemispheric asymmetry were found to be the contrast and the visual field size. The neuropsychological research on brain-injured patients has given some further evidence of the hemispheric asymmetry in spatial frequency processing. In conclusion, it is argued that the major merit of the spatial frequency hypothesis was in the attempt to investigate the hemispheric specialization of lower and higher levels of visual information processing from the perspective of a unified computational conception of visual perception.

Brain↗

An fMRI study of sex differences in regional activation to a verbal and a spatial task.

Sex differences in cognitive performance have been documented, women performing better on some phonological tasks and men on spatial tasks. An earlier fMRI study suggested sex differences in distributed brain activation during phonological processing, with bilateral activation seen in women while men showed primarily left-lateralized activation. This blood oxygen level-dependent fMRI study examined sex differences (14 men, 13 women) in activation for a spatial task (judgment of line orientation) compared to a verbal-reasoning task (analogies) that does not typically show sex differences. Task difficulty was manipulated. Hypothesized ROI-based analysis documented the expected left-lateralized changes for the verbal task in the inferior parietal and planum temporal regions in both men and women, but only men showed right-lateralized increase for the spatial task in these regions. Image-based analysis revealed a distributed network of cortical regions activated by the tasks, which consisted of the lateral frontal, medial frontal, mid-temporal, occipitoparietal, and occipital regions. The activation was more left lateralized for the verbal and more right for the spatial tasks, but men also showed some left activation for the spatial task, which was not seen in women. Increased task difficulty produced more distributed activation for the verbal and more circumscribed activation for the spatial task. The results suggest that failure to activate the appropriate hemisphere in regions directly involved in task performance may explain certain sex differences in performance. They also extend, for a spatial task, the principle that bilateral activation in a distributed cognitive system underlies sex differences in performance.

Brain↗

Neuroanatomic overlap of working memory and spatial attention networks: a functional MRI comparison within subjects.

Frontal and posterior parietal activations have been reported in numerous studies of working memory and visuospatial attention. To directly compare the brain regions engaged by these two cognitive functions, the same set of subjects consecutively participated in tasks of working memory and spatial attention while undergoing functional MRI (fMRI). The working memory task required the subject to maintain an on-line representation of foveally displayed letters against a background of distracters. The spatial attention task required the subject to shift visual attention covertly in response to a centrally presented directional cue. The spatial attention task had no working memory requirement, and the working memory task had no covert spatial attention requirement. Subjects' ability to maintain central fixation was confirmed outside the MRI scanner using infrared oculography. According to cognitive conjunction analysis, the set of activations common to both tasks included the intraparietal sulcus, ventral precentral sulcus, supplementary motor area, frontal eye fields, thalamus, cerebellum, left temporal neocortex, and right insula. Double-subtraction analyses yielded additional activations attributable to verbal working memory in premotor cortex, left inferior prefrontal cortex, right inferior parietal lobule, precuneus, and right cerebellum. Additional activations attributable to covert spatial attention included the occipitotemporal junction and extrastriate cortex. The use of two different tasks in the same set of subjects allowed us to provide an unequivocal demonstration that the neural networks subserving spatial attention and working memory intersect at several frontoparietal sites. These findings support the view that major cognitive domains are represented by partially overlapping large-scale neural networks. The presence of this overlap also suggests that spatial attention and working memory share common cognitive features related to the dynamic shifting of attentional resources.

Adult↗

Neural correlates of naming actions and of naming spatial relations.

Ina [(15)O] water PET experiment, 10 normal subjects retrieved words denoting actions (performed with or without an implement), and another 10 normal subjects retrieved words denoting the spatial relations between objects. Our objective was to test the following hypothesis: that the salient neural activity associated with naming actions and spatial relations occurs in left frontal operculum and left parietal association cortices, but not in the left inferotemporal cortices (IT) or in the right parietal association cortices. There were two control tasks, one requiring a decision on the orientation of unknown faces (a standard control task in our laboratory) and another requiring the retrieval of words denoting the concrete entities used in the action and spatial relations tasks. In accordance with the hypothesis, both naming actions and spatial relations (using the face orientation task as control activated the left frontal operculum; naming actions also activated the left parietal lobe. However, sectors of the left posterior IT were also engaged in both naming actions and spatial relations. When the naming of concrete entities was subtracted from the naming of actions performed with such entities, area MT in the posterior temporo-occipital region was activated bilaterally. On the other hand, when naming of the concrete entities was subtracted from the naming of spatial relations, left parietal activation was found, and when two tasks of naming spatial relations were contrasted to each other bilateral parietal activation was seen, right when abstract stimuli were used and left when concrete objects were used. The activity in posterior IT is thought to be related to object processing and possibly name retrieval at a subconscious level.

Adult↗

Spatial tuning of the pattern ERG across temporal frequency.

The spatial response function of the electroretinogram (ERG) to contrast checkerboard pattern reversal at several check sizes was determined at a fixed contrast. The influence of the rate of modulation on the spatial response function was assessed. Reversing square wave patterns were presented at eight temporal frequencies ranging from 1 to 25 reversals per sec. The waveform consisted of an initial positive and a subsequent negative deflection. Irrespective of the temporal frequency, the spatial response function of the positive component did not show a spatial tuning. The amplitude of the negative component exhibited a pronounced attenuation of the response at check sizes larger than optimal. Mean maximal amplitude was found at an optimal check size between 25 and 50 min of arc. A distinction between a positive or negative component was not made for temporal frequencies higher than 10 reversals per sec, since the waveform at these modulation rates consisted merely of a sinusoidal steady-state response. The spatial response function obtained at 14 reversals per sec, resembling that of the negative component, exhibited a prominent spatial tuning. The results demonstrate that the pattern ERG has at least two components: a positive component which is not specific to changes in retinal distribution of contrast, followed by a negative wave showing spatial tuning across temporal frequency.

Electroretinography↗

Distortion of spatial selectivity by pattern onset stimulation.

Spatial selectivity of pattern evoked potentials has been thought to provide evidence of lateral inhibition. However, spatial tuning functions may be distorted by pattern onset stimulation that is applied repeatedly to the same area of retina so an after-image is formed. This only applies at low spatial frequencies because of the randomizing effects of eye movements. Low-frequency attenuation may therefore be exaggerated. Pattern reversal stimulation has the opposite effect and this is reflected in the literature by fewer reports of bandpass functions. A new method has therefore been devised to provide the true spatial response function. The spatial phase is reversed after every two consecutive presentations. By combining this paradigm with a correction for the optical transfer function of the eye, the true neural response function is obtained. Ten subjects participated in this study to evaluate the distortion of spatial selectivity in the pattern electroretinogram. The new stimulus paradigm reduced the low spatial frequency attenuation to a barely significant level giving an almost flat amplitude response for the + ve and - ve transients of the pattern electroretinogram for check sizes from 222' to 7' angular subtense. However, correction for optical degradation produces bandpass curves, which closely correspond to those predicted from recent data on receptive fields of primate retinal ganglion cells.

Adult↗