The inversion of sensory processing by feedback pathways: a model of visual cognitive functions.

The mammalian visual system has a hierarchic structure with extensive reciprocal connections. A model is proposed in which the feedback pathways serve to modify afferent sensory stimuli in ways that enhance and complete sensory input patterns, suppress irrelevant features, and generate quasi-sensory patterns when afferent stimulation is weak or absent. Such inversion of sensory coding and feature extraction can be achieved by optimization processes in which scalar responses derived from high-level neural analyzers are used as cost functions to modify the filter properties of more peripheral sensory relays. An optimization algorithm, Alopex, which is used in the model, is readily implemented with known neural circuitry. The functioning of the system is investigated by computer simulations.

Multisensory integration: central processing modifies peripheral systems.

Multisensory integration in humans is thought to be essentially a brain phenomenon, but theories are silent as to the possible involvement of the peripheral nervous system. We provide evidence that this approach is insufficient. We report novel tactile-auditory and tactile-visual interactions in humans, demonstrating that a facilitating sound or visual stimulus that is exactly synchronous with an excitatory tactile signal presented at the lower leg increases the peripheral representation of that excitatory signal. These results demonstrate that during multisensory integration, the brain not only continuously binds information obtained from the senses, but also acts directly on that information by modulating activity at peripheral levels. We also discuss a theoretical framework to explain this novel interaction.

Insular cortex and neuropsychiatric disorders: a review of recent literature.

The insular cortex is located in the centre of the cerebral hemisphere, having connections with the primary and secondary somatosensory areas, anterior cingulate cortex, amygdaloid body, prefrontal cortex, superior temporal gyrus, temporal pole, orbitofrontal cortex, frontal and parietal opercula, primary and association auditory cortices, visual association cortex, olfactory bulb, hippocampus, entorhinal cortex, and motor cortex. Accordingly, dense connections exist among insular cortex neurons. The insular cortex is involved in the processing of visceral sensory, visceral motor, vestibular, attention, pain, emotion, verbal, motor information, inputs related to music and eating, in addition to gustatory, olfactory, visual, auditory, and tactile data. In this article, the literature on the relationship between the insular cortex and neuropsychiatric disorders was summarized following a computer search of the Pub-Med database. Recent neuroimaging data, including voxel based morphometry, PET and fMRI, revealed that the insular cortex was involved in various neuropsychiatric diseases such as mood disorders, panic disorders, PTSD, obsessive-compulsive disorders, eating disorders, and schizophrenia. Investigations of functions and connections of the insular cortex suggest that sensory information including gustatory, olfactory, visual, auditory, and tactile inputs converge on the insular cortex, and that these multimodal sensory information may be integrated there.

Monitors: key mechanisms and roles in the development and aging of the consciousness and self.

A network of interacting neural structures, called monitors, exists in the mammalian brain in which data derived from sensory inputs and from memory stores is precisely displayed within the brain. The key function of monitors is to provide an 'ultimate monitor', proposed to be the locus that generates the phenomenon of conscious self awareness, with information that defines or maps the positions of the parts of an individual with respect to each other and with respect to external objects or events at specific times. The resolution of at least some of these monitors (e.g. some concerned with vision) is extremely great and approaches, in the case of vision, the precision with which images of external objects are projected onto the retina. This conclusion is based on the fact that an individual is able to perceive visual images with an acuity that closely approximates the fineness of resolution of the retinal image. The sensory signals that provide information about body part positions and those that provide information about the exterior are evidently integrated with each other in a suitable hierarchy of monitors so as to provide a coherent representation of self-vs.-environment. The logical 'framework' monitor for this integrated display-mapping is proposed to be that that maps the body in space and it is proposed that the locations of objects perceived through the touch sense and senses that deal with more remote items in the environment become superimposed on a map that extends or extrapolates the body space map beyond the body's physical boundaries, a learning process that occurs during development. The ultimate monitor not only receives a display of the synthetic representations derived currently through the integrative functions defined above, but also is provided with at least four other inputs from other different classes of monitors. One of these is a monitoring system that generates timing signals needed to separate inputs into a time order and to assign an order to them. It is proposed that it is awareness of these timing signals by the ultimate monitor that is the essential and indispensible input that generates the phenomenon of awareness. A second input to the monitor that is the self is a selected part of its own activities. This awareness of what the ultimate monitor is receiving, doing or planning to do in the future is the characteristic necessary for awareness of self.(ABSTRACT TRUNCATED AT 400 WORDS)

Intensity coding of auditory stimuli: an fMRI study.

The effect of stimulus intensity (sound pressure level, SPL) of auditory stimuli on the BOLD response in the auditory cortex was investigated in 14 young and healthy subjects, with no hearing abnormalities, using echo-planar, functional magnetic resonance imaging (fMRI) during a verbal and a non-verbal auditory discrimination task. The stimuli were presented block-wise at three different intensities: 95, 85 and 75 dB (SPL). All subjects showed fMRI signal increases in superior temporal gyrus (STG) covering primary and secondary auditory cortex. Most importantly, the spatial extent of the fMRI response in STG increased with increasing stimulus intensity. It is hypothesized that spreading of excitation is associated with the encoding of increasing stimulus intensity levels. In addition, we found bifrontal activation supposedly evoked by the auditory-articulary loop of working memory. The results presented here should assist in the design of optimal activation strategies for studying the auditory cortex with fMRI paradigms and may help in understanding intensity coding of auditory stimuli.

Spiking Phineas Gage: a neurocomputational theory of cognitive-affective integration in decision making.

The authors present a neurological theory of how cognitive information and emotional information are integrated in the nucleus accumbens during effective decision making. They describe how the nucleus accumbens acts as a gateway to integrate cognitive information from the ventromedial prefrontal cortex and the hippocampus with emotional information from the amygdala. The authors have modeled this integration by a network of spiking artificial neurons organized into separate areas and used this computational model to simulate 2 kinds of cognitive-affective integration. The model simulates successful performance by people with normal cognitive-affective integration. The model also simulates the historical case of Phineas Gage as well as subsequent patients whose ability to make decisions became impeded by damage to the ventromedial prefrontal cortex.

Intensity discrimination and auditory brainstem responses in cochlear implant and normal-hearing listeners.

Intensity-discrimination limens (IDLs) and auditory brainstem responses (ABRs) were measured as a function of stimulus intensity in 6 cochlear implant (CI) and 8 normal-hearing (NH) listeners. Pulse-train stimuli were delivered electrically to the auditory nerve in CI listeners and acoustically in NH listeners. In CI listeners, the IDLs expressed as Weber fractions decreased monotonically with increasing intensity. In NH listeners, a nonmonotonic IDL function showing a peak a midintensities was observed. ABR wave amplitudes increased regularly with intensity only in CI listeners. Results support the notion that the slight decrease in Weber's fractions with increasing sound intensity--generally referred to as "the near-miss to Weber's law"--is subtended by retrocochlear processes, whereas the increase in Weber's fractions at midlevels--known as "the severe departure from Weber's law"--originates in cochlear mechanisms.

Contributions of the hippocampus, amygdala, and dorsal striatum to the response elicited by reward reduction.

Rats were trained to run down a runway for either 1 or 10 food pellets. After training, those receiving 10 pellets were shifted to 1 pellet. Such shifts typically elicit a temporary decrease in running speed. Groups of normal rats and rats with bilateral lesions of the fimbria-fornix, lateral-basolateral complex of the amygdala, or dorsal striatum were tested with the shifted and unshifted procedures. Separate experiments, identical except for the intertrial intervals (ITIs; 3 min vs. 30 s), were carried out. The data are consistent with the view that an integrated action of multiple neural systems is required to observe the typical response to reward reduction in unlesioned rats. One system that includes the dorsal striatum promotes a reinforced approach response to the goal box. A neural system that includes fimbria-fornix is required to retain information about reduced reward over the 3-min ITI. A system that includes the amygdala may acquire a conditioned aversive response to the goal box after the shift is detected, leading to reduced speeds over testing.

Comparison of ventral subicular and hippocampal neuron spatial firing patterns in complex and simplified environments.

Activity from ventral subicular and hippocampal CA1 neurons was recorded in rats exploring a 4-arm radial maze in which the local and distal cues could be manipulated. Cells from both regions exhibited place fields, although ventral subicular neurons had larger fields than hippocampal cells. Rotation of the local and distal cues in opposite directions produced movement of the place fields in either direction or a complete change in firing pattern. Simplifying the environment also produced changes in place field location. Despite similarities between regions, subiculum fields decreased in size whereas hippocampal fields increased in the simple environment. These findings suggest that subicular cells may receive converging input from several hippocampal neurons and code more complex configurations of the cues.

Cue control and head direction cells.

Previous research has shown that head direction (HD) cells in both the anterior dorsal thalamus (ADN) and the postsubiculum (PoS) in rats discharge in relation to familiar, visual landmarks in the environment. This study assessed whether PoS and ADN HD cells would be similarly responsive to nonvisual or unfamiliar environmental cues. After visual input was eliminated by blindfolding the rats, HD cells maintained direction-specific discharge, but their preferred firing directions became less stable. In addition, rotations of the behavioral apparatus indicated that some nonvisual cues (presumably tactile, olfactory, or both) exerted above chance stimulus control over a cell's preferred firing direction. However, a prominent auditory cue was not effective in exerting stimulus control over a cell's preferred direction. HD cell activity also was assessed after rotation of a novel visual cue exposed to the rat for 1, 3, or 8 min. An 8-min exposure was enough time for a novel visual cue to gain control over a cell's preferred direction, whereas an exposure of 1 or 3 min led to control in only about half the sessions. These latter results indicate that HD cells rely on a rapid learning mechanism to develop associations with landmark cues.

Taste responses in the greater superficial petrosal nerve: substantial sodium salt and amiloride sensitivities demonstrated in two rat strains.

A great quantity of research has focused on neural responses of the chorda tympani nerve (CT) to taste stimuli. This report examined salt and sugar sensitivity of the greater superficial petrosal nerve (GSP) and the effect of amiloride on these neural responses. In addition to Sprague-Dawley (SD) rats that have CT responses typical of most rat strains, we included Fischer 344 (F344) rats whose CT responses to sodium chloride (NaCl) are higher than those of other strains. After a stimulation series in which water served as the rinse, a series of stimuli was presented in 100 microM amiloride. The GSP was highly responsive to NaCl, sodium acetate (NaAc), ammonium chloride, and sucrose; NaCl and NaAc responses were strongly suppressed by amiloride. Relative responses to NaCl were significantly higher in F344 than in SD rats. In summary, the GSP is highly sensitive to salt and sugar stimulation, and palatal taste receptors have a considerable degree of amiloride sensitivity.

On the specificity of neurons and visual areas.

The dominant view during the past 40 years has been that the visual system analyzes the visual scene by breaking it down into basic attributes such as color, form, motion, depth and texture. Individual dedicated neurons and specific visual areas were believed to be devoted to the analysis of each of these attributes. Current research has challenged these views by emphasizing that neurons, especially in the cortex, have multifunctional properties and therefore serve as general-purpose analyzers rather than feature detectors. Consequently, it appears that most extrastriate visual areas, rather than each being devoted to the analysis of a specific basic visual attribute, perform several different tasks and thereby engage in more advanced and complex analyses than had been realized.

Cortical mechanisms for visual perception of object motion and position in space.

The present review is aimed at analyzing and discussing some of the cortical mechanisms possibly involved in the perception of object motion and object localization in the visual field. A comprehensive approach to these topics would be beyond the scope of this work. The highest priority, therefore, will be given to the cortical machinery involved in these processes, while very little (or nothing at all) will be said on the possible role played by subcortical structures such as the lateral geniculate nucleus and the superior colliculus which, albeit not directly involved in perception, might contribute to it.

Functional streams in occipito-frontal connections in the monkey.

It is known that the prestriate cortical regions that project to area LIP in parietal cortex and to areas TEO and TE in temporal cortex are mostly separated. Two separate streams of information transfer from occipital cortex can this be distinguished. We wished to determine whether the parietal and temporal streams remain segregated in their projections to frontal cortex. Paired injections of retrograde fluorescent tracers were placed in parietal and temporal cortex, or in the lateral and medial parts of the frontal eye field (FEF). The cortical regions containing retrogradely labeled cells were reconstructed in two-dimensional maps. The results show that temporal cortex mainly projects to lateral FEF (area 45). Parietal cortex sends projections to medial FEF (area 8a) and to lateral FEF, as well as to area 46. Thus, the parietal and temporal streams converge in lateral FEF. Most of the occipital regions projecting to medial FEF are the same as those projecting to parietal cortex, whereas lateral FEF receives afferents from the same occipital regions as those sending projections to temporal cortex. Thus, one can distinguish two interconnected networks. One is associated with the inferotemporal cortex and includes areas of the ventral bank and fundus of the superior temporal sulcus (STS), lateral FEF and ventral prestriate cortex. This network emphasizes central vision, small accades and form recognition. The other network is linked to cortex of the intraparietal sulcus. It consists of areas of the upper bank and fundus of STS, medial FEF and dorsal prestriate cortex. These areas encode peripheral visual field and are active during large saccades.

A consideration of sensory factors involved in motor functions of the basal ganglia.

There is a sizeable literature concerning basal ganglia (BG) functioning that is based on data from experiments employing a method of analysis that is traditionally used with other motor areas. A brief review of this literature is presented and the following conclusion is reached: as compared to the success of traditional methodologies in elucidating the workings of other motor systems, their use in BG investigations has proven disappointing. A possible reason for the shortcomings of traditional analyses in BG research is discussed. The remainder of this review concerns an alternative approach to the study of the BG that follows from consideration of a variety of clinical and experimental findings. The literature suggests that sensory aspects of BG functioning must be taken into account to fully appreciate the role of this system in motor control. A review of the literature concerning the latter suggests two points: The BG function as sensory analyzer for motor systems. That is, the BG convert sensory data from a form that is receptor oriented to a form that is relevant for guiding movement. The BG ultimately affect movement by gating sensory inputs into other motor areas rather than by directly affecting these areas. This sensory-based model of BG functioning explains a number of apparent discrepancies in the literature. In addition, seemingly anomalous findings are reconciled with the overwhelming evidence that the BG are a motor system. In particular, the suggestions of a BG role in attention and cognition are viewed as being intrinsic rather than orthogonal to the role of the BG in movement.

Visual hemifield mapping using transcranial magnetic stimulation coregistered with cortical surfaces derived from magnetic resonance images.

The perception of a visual stimulus can be inhibited by occipital transcranial magnetic stimulation. This visual suppression effect has been attributed to disruption in the cortical gray matter of primary visual cortex or in the fiber tracts leading to V1 from the thalamus. However, others have suggested that the visual suppression effect is caused by disruption in secondary visual cortex. Here the authors used a figure-eight coil, which produces a focal magnetic field, and a Quadropulse stimulator to produce visual suppression contralateral to the stimulated hemisphere in five normal volunteer subjects. The authors coregistered the stimulation sites with magnetic resonance images in these same subjects using optical digitization. The stimulation sites were mapped onto the surface of the occipital lobes in three-dimensional reconstructions of the cortical surface to show the distribution of the visual suppression effect. The results were consistent with disruption of secondary visual cortical areas.

Spontaneous eye movements in goldfish: oculomotor integrator performance, plasticity, and dependence on visual feedback.

To quantify performance of the goldfish oculomotor neural integrator and determine its dependence on visual feedback, we measured the relationship between eye drift-velocity and position during spontaneous gaze fixations in the light and in the dark. In the light, drift-velocities were typically less than 1 deg/s, similar to those observed in humans. During brief periods in darkness, drift-velocities were only slightly larger, but showed greater variance. One hour in darkness degraded fixation-holding performance. These findings suggest that while visual feedback is not essential for online fixation stability, it may be used to tune the mechanism of persistent neural activity in the oculomotor integrator.

The effects of magnitude and direction of stimulus change on auditory event-related potentials elicited by deviant signal stimuli.

The present study sought to identify components of the auditory event-related potential (ERP) elicited by stimuli that serve as signals for overt discriminative responses. Sokolov's view of a selective neural filter for deviant stimuli predicts that responses to deviant signal stimuli will be graded in proportion to the amount of change from the standard and independent of the direction of that change. The demonstration of a bi-directional and graded ERP response requires at least two levels of stimulus change in each direction. The present study incorporated two deviants that were lower in pitch (lowest, low) and two that were higher in pitch (high, highest) in order to evaluate the degree (linear, quadratic, etc.) of the function relating ERP response to tonal deviance. Stimulus changes on both the direction and magnitude dimensions were also varied on a trial-by-trial, rather than on a block-by-block, basis which eliminated potential confounds with block or session differences, and discriminative responses were required to both standard and deviant tones, thereby investing both categories of stimuli with signal value. The amplitude of the P3 component associated with deviant stimuli showed close correspondence to the (quadratic) function predicted from the selective filter model. A late negative slow wave (NSW) at Fz and a positive slow wave (PSW) at Pz differentiated deviant tones from the standard but did not distinguish between the deviants themselves. A fronto-central NSW observed at Fz and Cz for initial standard tones was greater than the predominantly frontal NSW elicited by the deviant tones. The topographical differences in NSW elicited by the initial standard tone and by all deviant tones suggest that different processes are reflected in the NSW response to these stimuli.

Color selection and location selection in ERPs: differences, similarities and 'neural specificity'.

It was hypothesized that color selection consists of two stages. The first stage represents a feature specific selection in neural populations specialized in processing color. The second stage constitutes feature non-specific selections, related to executive attentional processes and/or motor processes. This hypothesis was tested by investigating the effects of selectively attending to a specific color, location, or conjunction of location and color on the ERPs elicited by briefly flashed gratings. The gratings differed on three dimensions: color (red or blue), location in the visual field (4.4 degrees to the left or right of fixation) and form (target or non-target). Subjects had to respond to the presentation of target gratings in the attended category. Color selection was reflected in an enhanced parietal positivity in the 150-190 ms interval. Source analyses suggested that this color selection positivity might be generated in the basal occipital cortex, possibly human V4, an area of the brain specialized in color processing. The effect was separated from the P1 spatial attention effect both in topography and sources. Color selection was also reflected in a contralateral occipitotemporal negativity, which resembled the N1 spatial attention effect both in timing and topography. And finally, color selection was reflected in an N2b component. This N2b was similar in timing, topography and sources to the N2b's elicited by location selection and conjunction selection. We suggested that the N2b reflects feature non-specific selection processes, elicited by a range of attended stimuli, and possibly reflects activity in the anterior cingulate cortex. The NP80 was unaffected by attention to color and/or location and localized in striate cortex.

Functional organization of the medial division of the medial geniculate body of the cat: tonotopic organization, spatial distribution of response properties and cortical connections.

The discharge properties of 735 single units located in the pars magnocellularis (M) of the medial division of the medial geniculate body (MGB) were studied in 23 nitrous oxide anesthetized cats in response to simple acoustic stimuli (clicks, noise and tone bursts). A systematic decrease of single unit characteristic frequencies (CF) was observed along electrode track portions crossing M from dorso-medial to ventro-lateral. These data indicate that M is tonotopically organized with an arrangement of low CF units latero-ventrally and high CF units dorso-medially. This preferential arrangement of single units as a function of their CF was consistent with the location and orientation of clusters of labeled cells in M resulting from wheat-germ agglutinin labeled with horseradish peroxidase (WGA-HRP) injections in CF defined loci in the anterior (AAF) or primary (AI) auditory cortical fields. The quality of the tonotopic arrangement was low caudally and increased in the rostral direction, indicating that this tonotopicity concerns mainly the anterior half of M. Response latencies to clicks, noise and tone bursts were on average longer in the posterior part of M than in its anterior part. Time-locking of discharges in response to repetitive acoustic pulses was more frequent anteriorly than posteriorly and the upper limiting rate of locking was on average higher rostrally (up to 200-300 Hz). In contrast, other response properties such as responsiveness to the various combinations of simple acoustic stimuli, response patterns and tuning were more randomly distributed in M, showing the whole range of response properties seen in the MGB. Data derived from several injections of WGA-HRP performed in distinct auditory cortical fields in several animals indicated that M projects to the tonotopic cortical fields (AAF, AI and PAF) as well as to the non-tonotopically organized secondary auditory cortex (AII). The contribution of M to the total thalamic input reaching each field of the auditory cortex was quantitatively more important for AAF (30%) and PAF (20%) than for AI and AII (about 10% each).