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PubMed2005

Functional imaging with cellular resolution reveals precise micro-architecture in visual cortex.

Neurons in the cerebral cortex are organized into anatomical columns, with ensembles of cells arranged from the surface to the white matter. Within a column, neurons often share functional properties, such as selectivity for stimulus orientation; columns with distinct properties, such as different preferred orientations, tile the cortical surface in orderly patterns. This functional architecture was discovered with the relatively sparse sampling of microelectrode recordings. Optical imaging of membrane voltage or metabolic activity elucidated the overall geometry of functional maps, but is averaged over many cells (resolution >100 microm). Consequently, the purity of functional domains and the precision of the borders between them could not be resolved. Here, we labelled thousands of neurons of the visual cortex with a calcium-sensitive indicator in vivo. We then imaged the activity of neuronal populations at single-cell resolution with two-photon microscopy up to a depth of 400 microm. In rat primary visual cortex, neurons had robust orientation selectivity but there was no discernible local structure; neighbouring neurons often responded to different orientations. In area 18 of cat visual cortex, functional maps were organized at a fine scale. Neurons with opposite preferences for stimulus direction were segregated with extraordinary spatial precision in three dimensions, with columnar borders one to two cells wide. These results indicate that cortical maps can be built with single-cell precision.

Analysis of Variance↗

PubMed2000

Neurotrophin-4/5 alters responses and blocks the effect of monocular deprivation in cat visual cortex during the critical period.

The mechanisms underlying changes in neural responses and connections in the visual cortex may be studied by occluding one eye during a critical period in early postnatal life. Under these conditions, neurons in the visual cortex rapidly lose their responses to the deprived eye and ultimately lose many of their inputs from that eye. Cats at the peak of the critical period received infusions of exogenous neurotrophin NT-4/5 into primary visual cortex beginning before a short period of monocular deprivation. Within areas affected by NT-4/5, cortical cells remained responsive to the deprived eye, and maps of ocular dominance were no longer evident using intrinsic-signal optical imaging. Cortical cells also became broadly tuned for stimulus orientation and less responsive to visual stimulation through either eye. These effects required at least 48 hr exposure to the neurotrophin and were specific for trkB, because they were not seen with the trkA or trkC ligands NGF or NT-3. Even after neurons had already lost their responses to the deprived eye, subsequent NT-4/5 infusion could restore them. The NT-4/5 effects were not seen after the critical period. Together, these results suggest that trkB activation during the critical period may promote promiscuous connections independent of correlated activity.

Aging↗

PubMed1987

Importance of the visual and vestibular cortex for self-motion perception in man (circularvection).

Circularvection (CV), the optokinetically induced perception of self-motion, is based neurophysiologically upon visual-vestibular convergence. It is yet not known which visual pathways--subcortical accessory optic tract and/or cortical striate projection--convey optokinetic information to the central vestibular system in order to make possible the convergence that has been found: the vestibular nuclei, the thalamus and the vestibular cortex. The functional significance of the visual cortex was demonstrated in 12 patients with homonymous hemianopia who neither perceived CV nor exhibited a postural destabilization when exposed to optokinetic pattern motion (yaw or roll) restricted to the scotoma. The functional significance of the vestibular cortex as well as ipsilateral visual-vestibular interaction was demonstrated in 4 (out of 20) patients with tumour lesions involving the vestibular cortex areas. They either failed to perceive CV or showed a significant increase of CV-latencies when monocular optokinetic stimulation was restricted to the ipsilateral visual cortex. Arguments for and against the following hypothesis are discussed: circularvection is induced by visual motion stimulation of the primary visual cortex which then activates vestibular nuclei neurons by descending pathways and which also informs the vestibular cortex that self-motion with a perceptual direction is involved. Determination of the velocity of CV is mediated by direct visual-vestibular cortex interaction, which most probably is also involved in the perceptual interpretation of motion perception: Self-motion versus object-motion.

Adult↗

PubMed1992

Development of synapses in macaque monkey striate cortex.

A quantitative electron-microscopic (EM) analysis of the development of synaptic density (number of synapses/100 microns2 neuropil) has been done in primary visual cortex (striate, area 17) of the Old World monkey Macaca nemestrina. A comparative EM morphological study of developing synaptic contacts also was done in the same tissue. We find that a few immature synaptic contacts are present at fetal (F) 75 days either in the marginal zone, which becomes layer 1, or in the deepest portion of the cortical plate, the future layer 6. At F90-140 days synaptic contacts are found throughout the cortical plate, but their density remains higher in lower cortical layers. By F140 days synaptic density averaged for all layers (10.9) is three times higher than at F90 days. Just before and after birth, synaptic density rises very rapidly to peak at postnatal (P) 12 weeks (63) and then declines slowly to reach adult values (37.7) between 2-6 years. This pattern was further tested by comparing synaptic density in layer 2 which contains the last cells generated in the striate cortex to that in layer 6 which contains the first cells generated in the striate cortex. Layer 6 contained the first synapses, and had a higher density up to F140 days (an "inside-to-outside" distribution). Synaptic density was equal in the two layers at F152 days and P2 days, but by P12 weeks synaptic density in layer 2 was 27% higher than that in layer 6 (an "outside-to-inside" distribution). After P12 weeks, the synaptic density declined 51% in layer 2 and 21% in layer 6 so that both layers achieved similar densities by P6 years. A light and EM comparison of neuropil and synaptic contact morphology finds that, at each age up to birth, synapses in layer 2 are generally less mature than those in layer 6, but these differences disappear shortly after birth. Between P6-24 weeks, synaptic contacts throughout the cortex acquire a mature morphology that clearly differentiates between asymmetric and symmetric types, although asymmetric contacts continue to acquire more postsynaptic density until adulthood. This complex developmental pattern suggests a sequence for synaptic developments which is more related to neuron birthdate than to the arrival of extrinsic pathways or developmental events occurring in specific laminae.

Animals↗

PubMed1999

A physiological correlate of the 'spotlight' of visual attention.

Here we identify a neural correlate of the ability to precisely direct visual attention to locations other than the center of gaze. Human subjects performed a task requiring shifts of visual attention (but not of gaze) from one location to the next within a dense array of targets and distracters while functional MRI was used to map corresponding displacements of neural activation within visual cortex. The cortical topography of the purely attention-driven activity precisely matched the topography of activity evoked by the cued targets when presented in isolation. Such retinotopic mapping of attention-related activation was found in primary visual cortex, as well as in dorsomedial and ventral occipital visual areas previously implicated in processing the attended target features. These results identify a physiological basis for the effects of spatially directed visual attention.

Adult↗

PubMed1999

Functional organization of surgically created visual circuits.

Lesions of cerebral targets of the retina in newborn hamsters, when combined with transection of lemniscal pathways to the primary auditory or somatosensory thalamic nuclei or the secondary thalamic visual nucleus, can induce the formation of permanent retinal projections to the deafferented non-visual structures. These projections are retinotopically organized and form functional synapses. Consequently, neurons in the auditory or somatosensory cortices, which normally are not driven by visual stimuli, become visually responsive and have receptive field properties that ressemble, in several important ways, those of neurons in the visual cortex of normal animals. The surgically-induced retinothalamo-cortical pathways can mediate visually guided behaviors whose normal substrate, the pathway from the retina to the primary visual cortex via the thalamic dorsal lateral geniculate nucleus, is missing.

Journal Article↗

PubMed1984

Synaptic characteristics of identified pyramidal and multipolar non-pyramidal neurons in the visual cortex of young and adult rabbits. A quantitative Golgi-electron microscope study.

The visual cortex of 20 day old rats and rabbits has been considered as mature on the basis of the observations that the dendritic arborization and the overall synaptic population have almost reached their adult stage in these animals. In the present study we have investigated the visual cortex of 20 day and 7 month old (adult) rabbits in order to determine whether this apparent adult appearance also holds for the synaptic organization of individual neurons. Neurons mainly located in layers III and IV of the primary visual cortex (area 17) were Golgi-impregnated, gold toned and deimpregnated and were then, after embedding in plastic, sectioned serially. The number and length of synaptic profiles, and the length of the neuronal boundaries were analysed in every tenth section. From these counts and measurements the size distribution of the synaptic discs, the number of synapses per 100 micron2 neuronal surface and the receptive surface expressed as the percentage of the total neuronal surface covered with synaptic contacts were estimated using stereological methods. At both ages studied, the density of synapses was significantly higher for the non-pyramidal neurons than for the pyramidal neurons. Differences in the amount of receptive surface were parallel to the differences observed for the number of synapses per 100 micron2. At day 20 the receptive surface of the non-pyramidal neurons was significantly larger than that of the pyramidal neurons. The receptive surface of the non-pyramidal neurons in the adult stage was not only larger than that of the pyramidal neurons in the adults, but also larger than that of the day 20 non-pyramidal neurons. From our results the following conclusions can be drawn: (1) The synaptic input received by the pyramidal neurons is mainly established at day 20 of postnatal life, i.e. prior to the establishment of adult visual behaviour. (2) The non-pyramidal neurons complete their maturation in a later stage than the pyramidal neurons. (3) Medium to large sized synaptic contacts are newly formed after day 20 and are mainly added to the synaptic population on dendrites of non-pyramidal neurons. (4) The specific increase in the number of synapses on non-pyramidal neurons is discussed in relation to intracortical inhibition which is thought to be important for the fine regulation of visual function during development.

Animals↗

PubMed1998

Spatial frequency processing in posteromedial lateral suprasylvian cortex does not depend on the projections from the striate-recipient zone of the cat's lateral posterior-pulvinar complex.

It is generally considered that the posteromedial part of the cat's lateral suprasylvian cortex is involved in the analysis of image motion. The main afferents of the posteromedial lateral suprasylvian cortex come from a direct retinogeniculate pathway and indirect retinotectal and retino-geniculo-cortical pathways. Removal of the primary visual cortex does not affect the spatial and temporal processing of suprasylvian cortex cells suggesting that these properties are derived from thalamic input. We have investigated the possibility that the striate-recipient zone of the lateral posterior nucleus-pulvinar complex may be responsible for the spatial (and temporal) frequency processing in posteromedial lateral suprasylvian cortex since these two regions establish strong bidirectional connections and share many visual properties. Experiments were done on anaesthetized normal adult cats. Visual responses in suprasylvian cortex were recorded before, during, and after the deactivation of the lateral part of the lateral posterior nucleus accomplished by the injection of lidocaine or GABA. Results can be summarized as follows. A total of 64 cells was tested. Out of this number, 11 units were affected by the deactivation of the lateral part of lateral posterior nucleus and one cell, by the blockade of pulvinar. For all cells, except one, the effect consisted in a global reduction of the evoked discharge rate suggesting that the thalamo-suprasylvian cortex projections are excitatory in nature. We did not find any significant differences in the optimal spatial frequency, nor in the width of the tuning function, whether the grating was presented at half- or saturation contrast. In addition, there were no significant differences between the low- and high cut-off spatial frequency values computed before and after the deactivation of the lateral posterior nucleus. No specific changes were observed in the contrast sensitivity function of the posteromedial lateral suprasylvian cortex cells. Similar results were observed with respect to the temporal frequency tuning functions. Deactivating the lateral posterior nucleus did not modify the direction selectivity nor the organization of the subregions of the lateral suprasylvian cortex "classical" receptive fields. The absence of strong changes in posteromedial lateral suprasylvian cortex cell response properties following the functional blockade of the lateral posterior nucleus suggests that the projections from this part of the thalamus are not essential to generate the spatial characteristics of most posteromedial lateral suprasylvian cortex receptive fields. These properties may be derived from other thalamic inputs (e.g., medial interlaminar nucleus) and/or from the intrinsic computation of the afferent signals within the lateral suprasylvian cortex. On the other hand, it is possible that the lateral posterior nucleus lateral suprasylvian cortex loop may be involved in other functions such as the analysis of complex motion as suggested by the findings from our and other groups.

Animals↗

PubMed1995

Blockade of action potential activity alters initial arborization of thalamic axons within cortical layer 4.

In the formation of connections during the development of the nervous system, it is generally accepted that there is an early phase not requiring neural activity and a later activity-dependent phase. The initial processes of axonal pathfinding and target selection are not thought to require neural activity, whereas the later fine-tuning of connections into their final adult patterns does. We report an apparent exception to this rule in which action potential activity seems to be required very early in development for thalamic axons to form appropriate patterns of terminal arborizations with their ultimate target neurons in layer 4 of the cerebral cortex. Blockade of sodium action potentials during the 2-week fetal period when visual thalamic axons initially grow into the primary visual cortex in cats prevents the normally occurring branching of lateral geniculate nucleus axons within layer 4. This observation implies a role for action-potential activity in cerebral cortical development far earlier than previously suspected, weeks before eye-opening and the onset of the well-known process of activity-dependent reorganization of axonal terminal arbors that leads to the formation of ocular dominance columns.

Action Potentials↗

PubMed2004

Different types of signal coupling in the visual cortex related to neural mechanisms of associative processing and perception.

The hypothesis of object representation by synchronization in the visual cortex has been supported by our recent experiments in monkeys. They demonstrated local synchrony among gamma activities (30-90 Hz) and their perceptual modulation, according to the rules of figure-ground segregation. However, gamma-synchrony in primary visual cortex is restricted to few mm, challenging the synchronization hypothesis for larger cortical object representations. The restriction is due to randomly changing phase relations among locally synchronized patches which, however, form continuous waves of gamma-activity, traveling across object representations. The phase continuity of these waves may support coding of object continuity. Interactions across still larger distances, measured among cortical areas in human data, involve amplitude envelopes of gamma signals. Based on models with spiking neurons we discuss potentially underlying mechanisms. Most important for gamma synchronization are local facilitatory connections with distance-dependent delays. They also explain the occurrence of gamma waves and the restriction of gamma-synchrony. Fast local feedback inhibition generates gamma oscillations and supports local synchrony, while slow shunting inhibitory feedback supports figure-ground segregation. Finally, dispersion in inter-areal far projections destroys coherence of gamma signals, but preserves their amplitude modulations. In conclusion, we propose that the hypothesis of associative processing by gamma synchronization be extended to more general forms of signal coupling.

Action Potentials↗

PubMed2004

Stimulus locking and feature selectivity prevail in complementary frequency ranges of V1 local field potentials.

The local field potential (LFP) is a population measure of neuronal activity complementary to spike trains. Whereas the response properties of the spiking activity in the visual cortex have been characterized extensively, the responses of the LFP have not been well explored. No coherent picture exists about which frequency ranges exhibit feature tuning or show stimulus locked activity. Addressing this, we recorded LFP in the primary visual cortex of alert cats and calculated the tuning indices for orientation, spatial and temporal frequency. Furthermore, we quantified the locking of the power in different LFP frequency bands to the velocity profile of artificial and natural stimuli. We found that the LFP in alert animals is well tuned with similar specificity to orientation, spatial frequency and temporal frequency. Tuning to these features is most prominent in two frequency bands (8-23 Hz and 39-109 Hz). In two complementary frequency bands (23-39 Hz and above 109 Hz) the dynamics of the LFP power is locked tightly to the temporal structure of the stimulus. This locking is furthermore independent of the spatial structure of the stimulus. Together these four frequency bands cover the whole frequency range investigated. In contrast to previous studies, which often reported correlates of visual processing only in a limited frequency range of the LFP, the present results suggest that the entire frequency range of the LFP can be assigned a role in visual processing.

Action Potentials↗

PubMed2006

The neuronal structure of the dorsal nucleus of the lateral geniculate body in the common shrew (Sorex araneus) and the bank vole (Clethrionomys glareolus): Golgi and Nissl studies.

The topography and neuronal structure of the dorsal nucleus of the lateral geniculate body (GLd) of the common shrew and the bank vole are similar. The lateral geniculate body of both the species examined has a homogeneous structure and no observable cytoarchitectonic lamination. On the basis of the shape of the dendritic arbours as well as the pattern of dendritic arborisations the following two types of neurons were distinguished. Type I "bushy" neurons that have multipolar or round perikarya (common shrew perikarya 9-12 microm, bank vole perikarya 10-13 microm), with 4-6 short thick dendritic trunks that subdivide into many bush-like branches. The dendritic trunks are smooth, in contrast to the distal branches, which are covered with numerous spine-like protrusions of different lengths and forms. An axon emerges from the soma, sometimes very close to one of the primary dendrites. The type I neurons are typically projection cells that send their axons to the primary visual cortex. These neurons predominate in the GLd of both species. Type II neurons, which have an elongated soma with primary dendrites arising from opposite poles of the perikaryon (common shrew perikarya 8-10 microm, bank vole perikarya 9-11 microm). The dendritic arbours of these cells are less extensive and their dendrites have fewer spines than those of the type I neurons. Axons were seldom observed. The type II neurons are presumably interneurons and are definitely less numerous than the type I neurons.

Animals↗

PubMed2003

1 Hz rTMS enhances extrastriate cortex activity in migraine: evidence of a reduced inhibition?

We recently reported a paradoxical facilitatory effect of 1 Hz repetitive TMS (rTMS) on the primary visual cortex in migraine possibly due to the failure of inhibitory circuits, unable to be upregulated by low frequency rTMS. To investigate if inhibitory circuit dysfunction extends beyond striate cortex in migraine with aura, we studied the effects of 1 Hz rTMS over the right extrastriate cortex on perception of illusory contours in these patients. Low-frequency rTMS enhanced activity of extrastriate cortex in migraineurs, speeding up reaction times on illusory contour perception. This finding supports the view of a failure of inhibitory circuits also involving the extrastriate cortex in migraine with aura.

Adult↗

PubMed1994

Correlational imaging of thalamocortical coupling in the primary visual pathway of the human brain.

While the anatomy of the human brain is well defined, the functional connectivity of its structures is far less understood. Modern neuroimaging techniques offer the unique opportunity of visualizing physiologic activation in central nervous structures and of identifying the elements underlying distributed networks for information processing. Following improved spatial resolution of deoxyhemoglobin-sensitive magnetic resonance imaging, we were able to detect simultaneous signal changes in the lateral geniculate nucleus and primary visual cortex during periodic photic stimulation. Visualization of coupled activation by cross-correlation analysis resulted in the first demonstration of thalamocortical interaction in the primary visual pathway of the intact human brain.

Adult↗

PubMed2006

Neuroimaging in human amblyopia.

Functional magnetic resonance imaging (fMRI), positron emission tomography (PET) and magnetoencephalography (MEG) have been the principal neuroimaging tools used to assess the site and nature of cortical deficits in human amblyopia. A review of this growing body of work is presented here with particular reference to various controversial issues, including whether or not the primary visual cortex is dysfunctional, the involvement of higher-order visual areas, neural differences between strabismic and anisometropic amblyopes, and the effects of modern-day drug treatments. We also present our own recent MEG work in which we used the analysis technique of synthetic aperture magnetometry (SAM) to examine the effects of strabismic amblyopia on cortical function. Our results provide evidence that the neuronal assembly associated with form perception in the extrastriate cortex may be dysfunctional in amblyopia, and that the nature of this dysfunction may relate to a change in the normal temporal pattern of neuronal discharges. Based on these results and existing literature, we conclude that a number of cortical areas show reduced levels of activation in amblyopia, including primary and secondary visual areas and regions within the parieto-occipital cortex and ventral temporal cortex.

Amblyopia↗

PubMed1992

Modification induced in visual cortical activity of rat by neonatal monocular enucleation.

Monocular enucleation was performed on rats at birth. The animals were raised and from the age of 3 months the evoked activity was tested in the contralateral visual cortex both by mapping of evoked potentials and by autoradiography. It was found that monocular enucleation changed the distribution of the evoked activity characteristically. The focus of activity shifted laterally and was restricted to the binocular part of the primary visual cortex, while hardly any evoked activity or labelled neurons were found in its medial part.

Animals↗

PubMed2005

Local networks in visual cortex and their influence on neuronal responses and dynamics.

Networks of neurons in the cerebral cortex generate complex outputs that are not simply predicted by their inputs. These emergent responses underlie the function of the cortex. Understanding how cortical networks carry out such transformations requires a description of the responses of individual neurons and of their networks at multiple levels of analysis. We focus on orientation selectivity in primary visual cortex as a model system to understand cortical network computations. Recent experiments in our laboratory and others provide significant insight into how cortical networks generate and maintain orientation selectivity. We first review evidence for the diversity of orientation tuning characteristics in visual cortex. We then describe experiments that combine optical imaging of orientation maps with intracellular and extracellular recordings from individual neurons at known locations in the orientation map. The data indicate that excitatory and inhibitory synaptic inputs are summed across the cortex in a manner that is consistent with simple rules of integration of local inputs. These rules arise from known anatomical projection patterns in visual cortex. We propose that the generation and plasticity of orientation tuning is strongly influenced by local cortical networks-the diversity of these properties arises in part from the diversity of neighbourhood features that derive from the orientation map.

Animals↗

PubMed2001

Neurons with two sites of synaptic integration learn invariant representations.

Neurons in mammalian cerebral cortex combine specific responses with respect to some stimulus features with invariant responses to other stimulus features. For example, in primary visual cortex, complex cells code for orientation of a contour but ignore its position to a certain degree. In higher areas, such as the inferotemporal cortex, translation-invariant, rotation-invariant, and even view point-invariant responses can be observed. Such properties are of obvious interest to artificial systems performing tasks like pattern recognition. It remains to be resolved how such response properties develop in biological systems. Here we present an unsupervised learning rule that addresses this problem. It is based on a neuron model with two sites of synaptic integration, allowing qualitatively different effects of input to basal and apical dendritic trees, respectively. Without supervision, the system learns to extract invariance properties using temporal or spatial continuity of stimuli. Furthermore, top-down information can be smoothly integrated in the same framework. Thus, this model lends a physiological implementation to approaches of unsupervised learning of invariant-response properties.

Algorithms↗