See me, hear me, touch me: multisensory integration in lateral occipital-temporal cortex

Our understanding of multisensory integration has advanced because of recent functional neuroimaging studies of three areas in human lateral occipito-temporal cortex: superior temporal sulcus, area LO and area MT (V5). Superior temporal sulcus is activated strongly in response to meaningful auditory and visual stimuli, but responses to tactile stimuli have not been well studied. Area LO shows strong activation in response to both visual and tactile shape information, but not to auditory representations of objects. Area MT, an important region for processing visual motion, also shows weak activation in response to tactile motion, and a signal that drops below resting baseline in response to auditory motion. Within superior temporal sulcus, a patchy organization of regions is activated in response to auditory, visual and multisensory stimuli. This organization appears similar to that observed in polysensory areas in macaque superior temporal sulcus, suggesting that it is an anatomical substrate for multisensory integration. A patchy organization might also be a neural mechanism for integrating disparate representations within individual sensory modalities, such as representations of visual form and visual motion.     

Extensive cochleotopic mapping of human auditory cortical fields obtained with phase-encoding FMRI

The primary sensory cortices are characterized by a topographical mapping of basic sensory features which is considered to deteriorate in higher-order areas in favor of complex sensory features. Recently, however, retinotopic maps were also discovered in the higher-order visual, parietal and prefrontal cortices. The discovery of these maps enabled the distinction between visual regions, clarified their function and hierarchical processing. Could such extension of topographical mapping to high-order processing regions apply to the auditory modality as well? This question has been studied previously in animal models but only sporadically in humans, whose anatomical and functional organization may differ from that of animals (e.g. unique verbal functions and Heschl''s gyrus curvature). Here we applied fMRI spectral analysis to investigate the cochleotopic organization of the human cerebral cortex. We found multiple mirror-symmetric novel cochleotopic maps covering most of the core and high-order human auditory cortex, including regions considered non-cochleotopic, stretching all the way to the superior temporal sulcus. These maps suggest that topographical mapping persists well beyond the auditory core and belt, and that the mirror-symmetry of topographical preferences may be a fundamental principle across sensory modalities.     

Evaluation of the validity of a maximum likelihood adaptive staircase procedure for measurement of olfactory detection threshold in mice

Threshold is defined as the stimulus intensity necessary for a subject to reach a specified percent correct on a detection test. MLPEST (maximum likelihood parameter estimation by sequential testing) is a method that is able to determine threshold accurately and more rapidly than many other methods. Originally developed for human auditory and visual tasks, it has been adapted for human olfactory and gustatory tests. In order to utilize this technique for olfactory testing in mice, we have adapted MLPEST methodology for use with computerized olfactometry as a tool to estimate odor detection thresholds. Here we present Monte Carlo simulations and operant conditioning data that demonstrate the potential utility of this technique in mice, we explore the ramifications of altering MLPEST test parameters on performance, and we discuss the advantages and disadvantages of using MLPEST compared to other methods for the estimation of thresholds in rodents. Using MLPEST, we find that olfactory detection thresholds in mice deficient for the cyclic nucleotide-gated channel subunit A2 are similar to those of wild-type animals for odorants the knockout animals are able to detect.     

Temporal dynamics of auditory and visual bistability reveal common principles of perceptual organization

When dealing with natural scenes, sensory systems have to process an often messy and ambiguous flow of information. A stable perceptual organization nevertheless has to be achieved in order to guide behavior. The neural mechanisms involved can be highlighted by intrinsically ambiguous situations. In such cases, bistable perception occurs: distinct interpretations of the unchanging stimulus alternate spontaneously in the mind of the observer. Bistable stimuli have been used extensively for more than two centuries to study visual perception. Here we demonstrate that bistable perception also occurs in the auditory modality. We compared the temporal dynamics of percept alternations observed during auditory streaming with those observed for visual plaids and the susceptibilities of both modalities to volitional control. Strong similarities indicate that auditory and visual alternations share common principles of perceptual bistability. The absence of correlation across modalities for subject-specific biases, however, suggests that these common principles are implemented at least partly independently across sensory modalities. We propose that visual and auditory perceptual organization could rely on distributed but functionally similar neural competition mechanisms aimed at resolving sensory ambiguities.     

哺乳动物大脑感觉皮层对多种感觉信息整合的研究进展

脑对多感觉信息的整合是人和高等动物获取环境中有意义信息的重要方式。长期以来科学界一直认为,脑对不同感觉刺激(包括视觉、听觉、躯体感觉等)信息的分析和加工由不同的感觉皮层介导,最终在联络皮层进行整合,形成综合性的感觉和意识,但最近的一些实验证据显示,以前被认为只负责对单一感觉刺激分析和处理的感觉皮层亦可受其他感觉刺激的影响并直接参与多感觉信息的整合作用,这些新的发现对过去传统的大脑皮层功能分区概念提出了严峻的挑战。就近些年来有关感觉皮层(主要包括听觉、视觉和躯体感觉皮层)对多感觉刺激信息整合的研究进行综述,以增加人们对大脑皮层功能的新认识,为感觉信息处理和编码及感觉信息整合的后续研究提供借鉴。     

A morphometric study of postnatal synaptogenesis in the neocortex of precocial and altricial rodents]

Quantitative electronmicroscopic studies have been made on the development of synapses in two modally different areas of the brain (V-VI layers of the visual and auditory cortex) in the rat and mouse Acomys cahirinus within first two weeks of their postnatal life. The density of synapses as well the relative amount of different types of synapses (symmetrical, asymmetrical, axo-spinal and synapses with large amounts of synaptic vesicles) were measured. It was shown that only in rats the development of synapses in the visual area usually is faster than in the auditory one.     

Perceptual organization of local elements into global shapes in the human visual cortex

The question of how local image features on the retina are integrated into perceived global shapes is central to our understanding of human visual perception. Psychophysical investigations have suggested that the emergence of a coherent visual percept, or a "good-Gestalt", is mediated by the perceptual organization of local features based on their similarity. However, the neural mechanisms that mediate unified shape perception in the human brain remain largely unknown. Using human fMRI, we demonstrate that not only higher occipitotemporal but also early retinotopic areas are involved in the perceptual organization and detection of global shapes. Specifically, these areas showed stronger fMRI responses to global contours consisting of collinear elements than to patterns of randomly oriented local elements. More importantly, decreased detection performance and fMRI activations were observed when misalignment of the contour elements disturbed the perceptual coherence of the contours. However, grouping of the misaligned contour elements by disparity resulted in increased performance and fMRI activations, suggesting that similar neural mechanisms may underlie grouping of local elements to global shapes by different visual features (orientation or disparity). Thus, these findings provide novel evidence for the role of both early feature integration processes and higher stages of visual analysis in coherent visual perception.     

The case for primate V3

The visual system in primates is represented by a remarkably large expanse of the cerebral cortex. While more precise investigative studies that can be performed in non-human primates contribute towards understanding the organization of the human brain, there are several issues of visual cortex organization in monkey species that remain unresolved. In all, more than 20 areas comprise the primate visual cortex, yet there is little agreement as to the exact number, size and visual field representation of all but three. A case in point is the third visual area, V3. It is found relatively early in the visual system hierarchy, yet over the last 40 years its organization and even its very existence have been a matter of debate among prominent neuroscientists. In this review, we discuss a large body of recent work that provides straightforward evidence for the existence of V3. In light of this, we then re-examine results from several seminal reports and provide parsimonious re-interpretations in favour of V3. We conclude with analysis of human and monkey functional magnetic resonance imaging literature to make the case that a complete V3 is an organizational feature of all primate species and may play a greater role in the dorsal stream of visual processing.     

Differences in the sensory support of the anterior and posterior sections of the limbic cortex of the rat

Using retrograde axonal transport of horseradish peroxidase, studies have been made on the thalamic projections in the anterior and posterior parts of the limbic cortex with special reference to exterosensory system projections (visual, auditory and somatic). Projections of the retinorecipient nuclei of the anterior hypothalamus and classic thalamic visual relays (n. geniculatus lateralis dorsalis, n. lateralis posterior, pretectum) were found in the anterior and posterior limbic cortex. There are also inputs from the thalamic relays of the auditory (n. geniculatus medialis) and somatic (n. ventralis posterior) systems in the posterior limbic cortex The data obtained indicate: 1) that sensory supply of the limbic cortex in rats may be realized via direct pathways from sensory thalamic relays; 2) that thalamic sensory supply of the anterior limbic cortex differs from that of the posterior one. In the former, projections of the thalamic relays of the visual, auditory and somatic systems were found, whereas in the posterior cortex only visual system is presented. Topographic organization of the thalamic nuclear areas sending afferents to the anterior limbic cortex differs from that of the posterior limbic cortex.     

Auditory cortex mapmaking: principles, projections, and plasticity

Maps of sensory receptor epithelia and computed features of the sensory environment are common elements of auditory, visual, and somatic sensory representations from the periphery to the cerebral cortex. Maps enhance the understanding of normal neural organization and its modification by pathology and experience. They underlie the derivation of the computational principles that govern sensory processing and the generation of perception. Despite their intuitive explanatory power, the functions of and rules for organizing maps and their plasticity are not well understood. Some puzzles of auditory cortical map organization are that few complete receptor maps are available and that even fewer computational maps are known beyond primary cortical areas. Neuroanatomical evidence suggests equally organized connectional patterns throughout the cortical hierarchy that might underlie map stability. Here, we consider the implications of auditory cortical map organization and its plasticity and evaluate the complementary role of maps in representation and computation from an auditory perspective.     

A polysensory pathway to the forebrain of the pigeon: the ascending projections of the nucleus dorsolateralis posterior thalami (DLP).

Visual evoked potentials (VEPs) in the associative neostriatum caudolaterale (NCL) have shorter latencies than those recorded in other visual forebrain areas. Therefore visual input into NCL probably stems from a subtelencephalic relay. Tracing experiments revealed a projection of the nucleus dorsolateralis posterior thalami (DLP) into those portions of NCL in which visual, auditory, and somatosensory afferents from intratelencephalic parasensory areas terminate. Since VEPs in NCL are abolished after DLP-lesions, this structure has to be the critical relay. However, DLP also projects to other associative forebrain areas and parts of the basal ganglia. Previous experiments had furthermore revealed that DLP-neurons integrate visual, auditory, and somatosensory inputs. Thus, the DLP-projection onto various associative forebrain areas represents a true polysensory thalamotelencephalic system.     

Hierarchy within human SI: supporting data from cytochrome oxidase,acetylcholinesterase and NADPH-diaphorase staining patterns

The human primary somatosensory cortex consists of four cytoarchitectonic subdivisions (3a, 3b, 1 and 2) that are likely to contain distinct somatosensory representations. The intraareal organization of these areas as well as that of the primary motor cortex (area 4) has been analyzed using histochemical stains of cytochrome oxidase, acetylcholinesterase and NADPH-diaphorase activity in normal human brains. Cytochrome oxidase activity was revealed in individual cortical neurons and neuropil. Areas 4, 3a and 3b were on average darker than areas 1 and 2. The laminar distribution of cytochrome oxidase activity varied in different areas. A prominent dark band was present in layers IV and lower III in areas 3a and 3b and in layer III in areas 1, 2 and 4. Acetylcholinesterase staining revealed fibers and pyramidal cells in layers III and V; stained layer III pyramids were rare in areas 3a and 3b and numerous in areas 1, 2 and 4. NADPH-diaphorase positive elements included Golgi-like stained non-pyramidal neurons and Nissl-like stained pyramidal neurons; the former were found, in small numbers, in layer II of areas 4, 3a, 3b and 1, and the latter in layers III and V of areas 4 and 3a and in layer V of areas 1 and 2. The dark cytochrome oxidase staining of layer IV and the paucity of acetylcholinesterase positive pyramids in areas 3a and 3b resemble the pattern found in primary visual and auditory areas, whereas the dark cytochrome oxidase staining in layer III and abundance of acetylcholinesterase positive pyramids in areas 1 and 2 that of association areas. These results suggest that the four areas included in human SI constitute hierarchical stages of cortical processing, with 3a and 3b corresponding to primary and 1 and 2 to secondary areas.     

Hierarchy within human SI: supporting data from cytochrome oxidase, acetylcholinesterase and NADPH-diaphorase staining patterns

The human primary somatosensory cortex consists of four cytoarchitectonic subdivisions (3a, 3b, 1 and 2) that are likely to contain distinct somatosensory representations. The intraareal organization of these areas as well as that of the primary motor cortex (area 4) has been analyzed using histochemical stains of cytochrome oxidase, acetylcholinesterase and NADPH-diaphorase activity in normal human brains. Cytochrome oxidase activity was revealed in individual cortical neurons and neuropil. Areas 4, 3a and 3b were on average darker than areas 1 and 2. The laminar distribution of cytochrome oxidase activity varied in different areas. A prominent dark band was present in layers IV and lower III in areas 3a and 3b and in layer III in areas 1, 2 and 4. Acetylcholinesterase staining revealed fibers and pyramidal cells in layers III and V; stained layer III pyramids were rare in areas 3a and 3b and numerous in areas 1, 2 and 4. NADPH-diaphorase positive elements included Golgi-like stained non-pyramidal neurons and Nissl-like stained pyramidal neurons; the former were found, in small numbers, in layer II of areas 4, 3a, 3b and 1, and the latter in layers III and V of areas 4 and 3a and in layer V of areas 1 and 2. The dark cytochrome oxidase staining of layer IV and the paucity of acetylcholinesterase positive pyramids in areas 3a and 3b resemble the pattern found in primary visual and auditory areas, whereas the dark cytochrome oxidase staining in layer III and abundance of acetylcholinesterase positive pyramids in areas 1 and 2 that of association areas. These results suggest that the four areas included in human SI constitute hierarchical stages of cortical processing, with 3a and 3b corresponding to primary and 1 and 2 to secondary areas.     

Linking visual perception with human brain activity.

The past year has seen great advances in the use of functional magnetic resonance imaging (fMRI) to study the functional organization of the human visual cortex, to measure the neuronal correlates of visual perception, and to test computational theories of vision. Activity in particular visual brain areas, as measured with fMRI, has been found to correlate with psychophysical performance, with visual attention, and with subjective perceptual experience.     

Prosimian sensory capacities

This paper reviews laboratory research on sensory capacities of prosimians. Most investigations to date have concentrated on visual and auditory sensitivity with relatively little attention directed toward olfactory and gustatory sensitivity. Diurnal prosimians have at least rudimentary color vision and appear deficient to anthropoids in other visual measures such as acuity. The most striking difference between prosimian and anthropoid auditory sensitivity is the increased sensitivity of the former group to high frequencies. No psychophysical data have been published on olfactory sensitivity of prosimians, but the relatively large olfactory areas in their brains and their propensity for scent marking strongly suggest that they have acute olfactory sensitivity. Most data on gustatory thresholds in prosimians does not permit sensitivity estimation since the data are joint functions of sensitivity as well as preferences.     

Functional regional asymmetry of the EEG intrahemispheric coherence in dogs during conditioning

Individual features of the regional interhemispheric relations in the brain were studied in dogs during alimentary conditioning. The electrical activity was recorded from symmetrical anterior (frontal and motor cortices) and posterior (visual and auditory cortices) areas of the neocortex. Comparison between the averaged left and right intrahemispheric EEG coherences revealed a dynamic character of interhemispheric relations dependent on the stage of conditioning. Individual features were shown. In a dog with strong type of the nervous system, in the anterior brain regions, the EEG coherence was higher in the left hemisphere than in the right one, whereas, on the contrary, in the posterior regions, the values were higher in the right than in the left hemisphere. In dogs with weak type of the nervous system, there was an inverse relationship. Thus, the spatial organization of the cortical electrical activity in the associative and projection brain areas was different.     

Processing local signals into global patterns

Perceptual organization or grouping is one of the central issues in vision research. Recent reports in the neuroimaging literature suggest that perceptual organization is mediated by distributed visual areas that range from the primary visual cortex (V1) to higher visual areas, depending on the availability of grouping cues and on the weight of contribution of each visual area. Evidence suggests that grouping by proximity and collinearity, and also perhaps filling-in, involve V1, whereas grouping by similarity and symmetry seems to depend on activation of higher visual areas. Further studies should include deliberate controls for confounding factors such as attentional artifacts and radial orientation bias, to clarify how spatiotemporal information in visual areas is integrated to give rise to perceptual organization.     

Functional architecture of auditory cortex

Three complementary approaches demonstrate new types of organization in rodent, feline and primate auditory cortex, as well as differences in processing between auditory and visual cortex. First, connectional work reveals patterns of thalamocortical and corticocortical input unique to the auditory cortex. Second, physiological studies find multiple, interleaved auditory processing modules related to corticocortical connections and embedded in the isofrequency gradient. Third, functional analyses demonstrate independent processing streams for sound localization and identification analogous to the 'what' and 'where' streams in visual cortex, although the modular arrangements are modality-specific. Taken together, these data show that the auditory cortex has common and unique functional substrates.     

Temporal Sequence of Visuo-Auditory Interaction in Multiple Areas of the Guinea Pig Visual Cortex

Recent studies in humans and monkeys have reported that acoustic stimulation influences visual responses in the primary visual cortex (V1). Such influences can be generated in V1, either by direct auditory projections or by feedback projections from extrastriate cortices. To test these hypotheses, cortical activities were recorded using optical imaging at a high spatiotemporal resolution from multiple areas of the guinea pig visual cortex, to visual and/or acoustic stimulations. Visuo-auditory interactions were evaluated according to differences between responses evoked by combined auditory and visual stimulation, and the sum of responses evoked by separate visual and auditory stimulations. Simultaneous presentation of visual and acoustic stimulations resulted in significant interactions in V1, which occurred earlier than in other visual areas. When acoustic stimulation preceded visual stimulation, significant visuo-auditory interactions were detected only in V1. These results suggest that V1 is a cortical origin of visuo-auditory interaction.     

Spatial organization of biopotentials and original features of visual images

Features of spatial organization of neocortical potentials during mental creating of the original and standard visual images were studied. Intrahemispheric and interhemispheric coherence in different EEG frequency bands and temporal relations between relative changes in the index of linear (correlation coefficient) and non-linear (multiple entropy) processes in different neocortical areas were analysed. Number of decreased coherence values in the high-frequency alpha subband between remote neocortical points during creating of an original image was significantly lower than during creating of a standard image. During creating of the original image, the number of synchronously functioning pairs of neocortical areas and the mean level of linear correlation between the areas were higher, and correlation by the non-linear index, on the contrary, was lower than during creation of the standard image. The correlation between changes in different neocortical areas for both processes during creating of the original image was only positive, and that during creating of the standard image was both positive and negative. The conclusion was made that creative and reproductive types of visual imagination were different in the spatial distribution of coordinated changes in the linear and non-linear processes, mean magnitudes of these changes, and the character of hemispheric interaction. The data on different interhemispheric relations in diagonal and central bilateral directions raise a question about the radial representation of visual imagination.