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Hippocampal mossy fibers and radial-maze learning in the mouse: a correlation with spatial working memory but not with non-spatial reference memory.

One hundred and eight male mice from nine different inbred strains were tested for two aspects of learning in an eight-arm radial maze. In the first experimental arrangement of the maze, measuring spatial working memory, clear strain differences were found on the fifth day of training. Furthermore, this type of learning showed a high positive correlation with the size of the intra- and infrapyramidal hippocampal mossy fiber terminal field as revealed with Timm's staining. In the second experiment, in which non-spatial reference memory was tested, significant strain differences were found for the learning variables, but there were no significant covariations with the sizes of the intra- and infrapyramidal mossy fiber terminal fields. These results, combined with previous data, suggest that heritable variations of the hippocampal intra- and infrapyramidal mossy fiber projection influence processes determining spatial learning capabilities in mice.

Animals↗

Spatial segregation between populations of ponto-cerebellar neurons: statistical analysis of multivariate spatial interactions.

This study applies terms and methods for describing spatial interactions between multivariate spatial point patterns, which are, to our knowledge, new in neurobiology. We consider two categories of points, type 1 and 2, distributed within a certain reference volume (such as a nucleus of the brainstem or a cortical area). The points may, for example, represent different categories of labelled cells or axonal fields of termination. We say that there is spatial neutrality between points of type 1 and 2 if the types are signed by random labelling. If a mechanism drives the two point categories together, we say that the point patterns are positively associated. Conversely, if a mechanism drives type 1 and 2 points apart, we say that they are segregated. By comparing two cumulative distribution functions of distances between points, we can distinguish neutrality, positive association, and segregation. One function, H12(t), is the cumulative distribution function of the distance t between a pair of randomly selected points of type 1 and 2. The other, H00(t), is the corresponding function for a pair of points randomly selected without reference to type. Plots of the estimated difference between these two functions give an indication of positive association, neutrality, or segregation. A statistical test, based on simulations of random (neutral) distributions, can be used to see whether deviations from neutrality are significant. We apply the analysis described above to a major pathway of the brain, namely the ponto-cerebellar projection. Different types of cells in the pontine nuclei are retrogradely labelled with the fluorescent tracers Rhodamine-B-isothiocyanate, Fluoro-Gold, and Fast Blue. The tracers are injected in adjacent or more distant folia of the cerebellar paraflocculus. The location of the somata of labelled cells are recorded and the total distribution reconstructed in three dimensions and displayed on a dynamic graphics workstation. We ask whether different units (folia) in the paraflocculus receive information from the same population, from two different positively associated populations, or from segregated cell populations. We find a statistically significant tendency for cell populations projecting to adjacent folia to be positively associated, although there are few cells containing multiple labels. Populations of neurons projecting to folia wider apart are significantly segregated. From inspections of the reconstructions, using real-time rotations, we find that the swarms of labelled neurons tend to accumulate in shells or lamellae in the pons. Within the lamellae, the cells are aggregated in clusters and bands with empty holes (containing unlabelled ponto-cerebellar cell bodies, presumably projecting to other cerebellar targets) in between.(ABSTRACT TRUNCATED AT 400 WORDS)

Amidines↗

The effects of large orientation and spatial frequency differences on spatial discriminations.

We have examined two questions: (1) can the finest orientation discrimination be achieved only between stimuli with similar spatial frequency content? and likewise, (2) can the lowest spatial frequency discrimination thresholds be achieved only with parallel gratings? In 2 AFC tests we found that neither type of discrimination was affected by stimulus differences along the other dimension. However, some small decreases in method of adjustment matching accuracy were associated with large differences along the secondary dimensions. Considering the neurophysiological implications, these data suggest that fine orientation and spatial frequency discrimination can occur even though separate populations of neurones in the primary visual cortex may be activated by the two stimuli to be discriminated.

Discrimination, Psychological↗

Spatial-frequency adaptation: evidence for a multiple-channel model of short-wavelength-sensitive-cone spatial vision.

The frequency selective effects of spatial adaptation were measured with vertically-oriented, cosine stimuli upon an intense long-wavelength (yellow) field, which isolated the short-wavelength-sensitive (S) cones. Consistent with isolated-S-cone spatial threshold and masking results, the adaptation measurements demonstrate S-cones input to multiple, orientation selective, spatial frequency mechanisms. Moreover, the adaptation measurements show the minimum number of S-cone mechanisms is three. The frequency tuning of each mechanism was derived from the S-cone threshold and masking results. Two of the tuning curves are bandpass with peak sensitivities in a vicinity of 0.7 and 1.4 c/deg, respectively. These two closely resemble tuning curves derived from results with luminance-modulated stimuli. Confined to the range of frequencies examined (0.25-2.83 c/deg), the third tuning curve is lowpass with a high-frequency cutoff of roughly 2.0 c/deg. However, subsequent measurements of orientation selectivity demonstrate the third mechanism to have bandpass frequency tuning as well.

Adaptation, Ocular↗

Phase-reversal discrimination in one and two dimensions: performance is limited by spatial repetition, not spatial frequency content.

Lawden [(1983) Vision Research, 23, 1451-1463] used vertical gratings containing two frequencies (F, nF) in phase discrimination (F + nF against F - nF) and compound detection (F + nF against F) experiments, where thresholds were measured by manipulating the contrast of the nF component. When n was varied, Lawden found a phase-plateau of moderate breadth where phase discrimination thresholds were about half of those measured in compound detection. I present the results of similar experiments, using one-dimensional (gratings) and two-dimensional (plaids). In a sine-plaid condition, the 1F grating was split into two 1F plaid components at +/- 45 deg from vertical while the nF component remained a vertical grating. In a square-wave plaid (SqW-plaid) condition the plaid components were square waves. For each of these conditions, the horizontal spatial repetition (SR) of the plaid is given by (F/square root of 2); it is half an octave lower than the spatial frequency (SF) of the oblique components but it is not represented in the stimulus spectrum. By plotting phase discrimination relative to compound detection a phase-plateau was found for all three conditions. When these data were plotted as a function of SF ratio (nF/F) the curves describing the two plaid conditions were found to be leftward translations of that describing the grating condition. However, when the results were plotted as a function of SR ratio (nF/SR), the three functions lay on top of each other. The finding that phase-reversal discrimination is not governed by the Fourier attributes of the stimulus per se, rules out an explanation in terms of a linear, broad-band, phase-sensitive mechanism. Rather, the results imply that information is combined across the set of SF- and orientation-tuned mechanisms before the decision variable. These interactions appear to be governed by the spatial (not Fourier) attributes of the luminance profile of the stimulus. A modified version of Bennett's [(1993) Perception & Psychophysics, 53, 292-304] phase discrimination model is presented as a post-hoc account of the data.

Contrast Sensitivity↗

Relationship between spatial integration and spatial spread of contrast energy in detection.

Detection efficiencies were measured for two kinds of grating stimuli. The stimuli of the first kind were uniform square shaped cosine gratings of various sizes but of constant spatial frequency. The stimuli of the second kind were composed of nine small grating patches of the same spatial frequency arranged into a square array. The array size was varied by changing the inter-patch distance. The efficiencies for the two kinds of stimuli obeyed the same decreasing function of area defined by the respective outlines of the grating patch array and the uniform grating. The result means that detection efficiency is not determined by the retinal area stimulated, but by the distances between different parts of stimulus, i.e. the spatial spread of contrast energy, which can be described by a radial moment measure computed from the image.

Contrast Sensitivity↗

The effects of hippocampal lesions upon spatial and non-spatial tests of working memory.

A series of experiments examined the proposal that the primary effect of hippocampal damage in rats is to disrupt working memory. Although extensive hippocampal lesions produced a severe impairment in forced-choice alternation--a test of spatial working memory--the same lesions did not impair the acquisition of a non-spatial test of working memory--delayed non-matching-to-sample. This test of object recognition required the rats to select that arm in a Y-maze which contained unfamiliar stimuli. Rats with hippocampal lesions were able to learn and perform this task at normal rates, even with retention delays of as long as 60 s. Two additional experiments helped confirm that the animals had indeed learnt a non-spatial test of working memory. The final experiment examined whether hippocampal lesions resulted in an increased sensitivity to proactive interference. It was found that repetition of test stimuli within a session, which increased interference, did attenuate recognition performance but there was no evidence that the animals with hippocampal lesions were differentially affected.

Animals↗

Role of the parahippocampal region in spatial and non-spatial memory: effects of parahippocampal lesions on rewarded alternation and concurrent object discrimination learning in the rat.

Rats with aspiration or excitotoxic (NMDA) lesions of the parahippocampal region were trained on a series of behavioral tasks which consisted of: (1) a test of spatial memory (discrete trial rewarded alternation), (2) a black-white discrimination, and (3) a test of non-spatial memory commonly used in primate models of amnesia (visual concurrent object discrimination). Rats in both lesion groups were severely impaired on the concurrent discrimination, even though they were able to learn the black-white discrimination normally. Animals with aspiration lesions were also impaired on the spatial memory task, whereas those with NMDA lesions did not differ from controls. The results indicate that concurrent object discrimination is a particularly sensitive measure of hippocampal/parahippocampal functions and suggest that these structures in the rat may serve mnemonic functions which are qualitatively similar to those of human and non-human primates.

Animals↗

The bailiwick of visuo-spatial working memory: evidence from unilateral spatial neglect.

An accurate representation of the visual environment is crucial for successful interaction with objects in that environment. The means by which that representation is formed in working memory is a major focus of the issues and research discussed in this paper. The discussion draws on experimental studies of healthy adults and of patients with impairments of visual perceptual processing or of visuo-spatial mental representation. These disorders are most commonly linked with the disorder referred to clinically as unilateral spatial neglect. We discuss the observed dissociation between perceptual neglect and representational neglect, and the phenomenon of implicit processing of information in the neglected hemifield. In so doing we explore the implications of the findings from this literature for the development of theories of visuo-spatial working memory.

Attention↗

Cerebral hemispheric specialization for spatial attention: spatial distribution of search-related eye fixations in the absence of neglect.

The "specialization" of the right hemisphere for spatial attention is widely accepted but poorly understood. While several theories have been supported by studies of patients with acute hemispatial neglect, generalizability beyond this population remains unclear. In this study, we compared the predictions of two attention models [Brain 119 (1996) 841; Trans. Am. Neurol. Assoc. 95 (1970) 143] when applied to data obtained from subjects with unilateral right- or left-cerebral lesions, but without clinical evidence of neglect during a visual search task. Both Left Lesion and Right Lesion subjects detected fewer targets in the contralesional hemispace. However, the Right Lesion subjects also made fewer visual fixations and longer saccades in the contralesional hemispace, suggesting a fundamental alteration in the architecture of visual search. The spatial distribution of fixations made by Right Lesion subjects more closely fits the prediction of a "salience" model than of the strict interpretation of a linear "gradient" model. These data support the long-standing notion of right hemisphere dominance for spatial attention, especially for the top-down processes entailed in self-directed visual search, and extend this to lesion patients without clinically evident neglect. A theoretical model based on the salience of extrapersonal space appears useful for understanding alterations of attentional allocation, particularly after recovery from stroke.

Adult↗

Spatial and non-spatial working memory at different stages of Parkinson's disease.

Groups of patients with Parkinson's disease, either medicated or unmedicated, were compared with a matched group of normal control subjects on a computerized battery of tests designed to assess spatial, verbal and visual working memory. In the spatial working memory task, subjects were required to search systematically through a number of boxes to find 'tokens' whilst avoiding those boxes in which tokens had previously been found. In the visual and verbal conditions, the subjects were required to search in exactly the same manner, but through a number of abstract designs or surnames, respectively, avoiding designs or names in which a token had previously been found. Medicated Parkinson's disease patients with severe clinical symptoms were impaired on all three tests of working memory. In contrast, medicated patients with mild clinical symptoms were impaired on the test of spatial working memory, but not on the verbal or visual working memory tasks. Non-medicated patients with mild clinical symptoms were unimpaired on all three tasks. These data are compared with the results of a previous study comparing groups of neurosurgical patients with frontal, temporal or amygdalo-hippocampectomy excisions on the same three tests of working memory. Taken together, the findings suggest that working memory deficits in Parkinson's disease emerge, and subsequently progress, according to a defined sequence, the evolution of which may be linked to the likely spatiotemporal progression of dopamine depletion within the striatum, in relation to the terminal distribution of its cortical afferents.

Antiparkinson Agents↗

Is second-order spatial loss in amblyopia explained by the loss of first-order spatial input?

The purpose of the study was to determine whether amblyopes show detection loss for second-order spatial information, and if present, whether the loss is explained by the loss of first-order spatial input. We psychophysically determined detection thresholds for the amblyopic and non-amblyopic eyes of five adult amblyopes and the dominant eyes of three control observers. We found that four amblyopic eyes and two non-amblyopic eyes showed second-order loss relative to the control eyes. The second-order loss was greater than the first-order loss at the carrier spatial frequency (first-order input). The extra second-order loss indicates an early amplification of cortical neural loss that we speculate is due to deficient binocular input to second-order neurons.

Adult↗

Bottom-up transfer of sensory-motor plasticity to recovery of spatial cognition: visuomotor adaptation and spatial neglect.

A large proportion of right-hemisphere stroke patients show hemispatial neglect, a neurological deficit of perception, attention, representation, and/or performing actions within their left-sided space, inducing many functional debilitating effects on everyday life, and responsible for poor functional recovery and ability to benefit from treatment. This spatial cognition disorder affects the orientation of behavior with a shift of proprioceptive representations toward the lesion side. This shift is similar to that produced by psychophysical manipulations as a wedge-prism exposure in normal healthy subjects. In both subjects, one major compensative effect of short-term prism adaptation is a shift of proprioceptive representations, demonstrated by a shift in manual straight-ahead pointing in the dark, in a direction opposite to the visual shift. In neglect patients, prism adaptation involves the shift of proprioceptive representations to the left with a reduction of rightward bias observed in neglect patients in visuo-manual tasks as line-bisection, line-cancellation or copy drawing. Improvement of neglect is also observed in no visuo-manual tasks as mental imagery, auditory extinction or posture. This generalization of prism adaptation effects at different neglect level symptoms suggests that the process of prism adaptation may activate brain functions related to multisensory integration and higher spatial representations. Moreover the positive effects found for both sensorimotor and more cognitive spatial functions lasted for at least two or more hours after prism removal. Unlike reduction of neglect through sensory stimulations, the long-lasting improvement of neglect after prism adaptation suggests the activation of short-term plasticity of brain functions related to coordinate transformations and space representations. Lastly, the duration of these effects could be useful in rehabilitation programs, as suggested by the effects of prism adaptation on disabling neglect symptoms as wheelchair driving, posture or writing.

Adaptation, Physiological↗

Colchicine-induced alterations of reference memory in rats: role of spatial versus non-spatial task components.

Male, Fischer-344 rats received bilateral injections of 2.5 micrograms of colchicine per site in the dorsal and ventral hippocampus. Intradentate colchicine preferentially destroyed dentate granule cells. Subsequent behavioral studies showed that 3 weeks after dosing, colchicine impaired the acquisition of a spatial, reference memory task in the Morris water maze. In a second group of rats, which were trained in the water maze prior to dosing, intradentate colchicine impaired retention of this task when rats were tested 3 weeks later. The acquisition of non-spatial reference memory task, an autoshape of a lever touch response in an operant chamber, was facilitated by prior administration of colchicine. Facilitative effects of colchicine were seen if delays of 0, 4, or 6 s were interposed between response and presentation of food reinforcement. If rats were trained to lever-touch with either a 0- or 4-s delay between response and reinforcement, intradentate colchicine had no effect on retention of the response 3 weeks later. These data are in accord with the conclusion that the dentate gyrus plays an important role in the acquisition of new information and retrieval of previously learned material and is an integral neural substrate for reference memory involving a spatial component.

Animals↗

Non-invasive detection of the single motor unit action potential by averaging the spatial potential distribution triggered on a spatially filtered motor unit action potential.

For research as well as diagnostic applications the non-invasive detection of the activity of single motor units is of interest. The most direct information is expected to be found in monopolarly recorded data. But when an array of surface electrodes is used for the monopolar recordings of the potential distribution on the skin, in most cases an additional invasive needle electrode is utilized to detect the exact points in time when a certain motor unit is firing. With this supplementary information, an averaging of the monopolar EMG tracings can be performed. In this paper, a completely non-invasive methodology is presented which replaces the invasive needle by a spatial filtering procedure. The EMG signals from the m. biceps brachii are recorded monopolarly with an electrode array. Afterwards, a spatial filtering procedure, called normal double differentiating filter, is applied to the data. The EMG signals obtained are investigated by means of an amplitude threshold to distinguish the activity of different motor units. The point of the maximum amplitude of the selected peaks then is used as trigger point to average the monopolar EMG data. The time courses of the motor unit action potential signals found after applying the described procedure show similar shapes, while two different components are to be identified: corresponding to the spread of the excitation, one is referring to stationary, the other to travelling events. These results justify the possibility to replace the needle electrode to obtain a trigger event in the future by the non-invasive spatial filtering procedure.

Action Potentials↗

Spatial and temporal contributions to the structure of spatial memory.

Three experiments investigated the effects of spatial and temporal contiguity in item recognition, location judgment, and distance estimation tasks. Ss learned the locations of object names in spatial arrays, which were divided into 2 regions. The names of locations were presented during map learning so that critical pairs appeared close in space and close in time, close in space but far in time, far in space but close in time, and far in space and far in time. Names primed each other in recognition only when they were neighbors in both space and time. In contrast, the effects of spatial and temporal contiguity in priming in location judgments were additive. Finally, temporal contiguity affected estimates of Euclidean distance when locations were close together, but not when they were far apart.

Adult↗

Spatial attention and implicit sequence learning: evidence for independent learning of spatial and nonspatial sequences.

This research investigated whether regular spatial orienting sequences can be learned implicitly and independently of response requirements. In a new version of a serial response task introduced by M. J. Nissen and P. Bullemer (1987) participants had to discriminate between objects that could occur at different locations. Independent sequences determined the succession of locations and objects. Even participants who were not aware of any regularities exhibited evidence for learning of both sequences (Experiment 1). Experiment 2 showed that the joint learning of spatial and object sequences was as efficient as learning of single sequences and that it even occurred when learning required memory for past sequence elements and attention was blocked through a secondary tone-counting task. Results are consistent with the idea that independent systems may exist for the implicit acquisition of spatial and nonspatial regularities.

Adult↗

Attention, spatial representation, and visual neglect: simulating emergent attention and spatial memory in the selective attention for identification model (SAIM).

The selective attention for identification model (SAIM) is presented. This uses a spatial window to select visual information for recognition, binding parts to objects and generating translation-invariant recognition. The model provides a qualitative account of both normal and disordered attention. Simulations of normal attention demonstrate 2-object costs and effects of object familiarity on selection, global precedence, spatial cueing, and inhibition of return. When lesioned, SAIM demonstrated either view- or object-centered neglect or spatial extinction, depending on the type and extent of lesion. The model provides a framework to unify (a) object- and space-based theories of normal selection, (b) dissociations within the syndrome of unilateral neglect, and (c) attentional and representational accounts of neglect.

Attention↗