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Dynamic properties of recurrent inhibition in primary visual cortex: contrast and orientation dependence of contextual effects.

A fundamental feature of neural circuitry in the primary visual cortex (V1) is the existence of recurrent excitatory connections between spiny neurons, recurrent inhibitory connections between smooth neurons, and local connections between excitatory and inhibitory neurons. We modeled the dynamic behavior of intermixed excitatory and inhibitory populations of cells in V1 that receive input from the classical receptive field (the receptive field center) through feedforward thalamocortical afferents, as well as input from outside the classical receptive field (the receptive field surround) via long-range intracortical connections. A counterintuitive result is that the response of oriented cells can be facilitated beyond optimal levels when the surround stimulus is cross-oriented with respect to the center and suppressed when the surround stimulus is iso-oriented. This effect is primarily due to changes in recurrent inhibition within a local circuit. Cross-oriented surround stimulation leads to a reduction of presynaptic inhibition and a supraoptimal response, whereas iso-oriented surround stimulation has the opposite effect. This mechanism is used to explain the orientation and contrast dependence of contextual interactions in primary visual cortex: responses to a center stimulus can be both strongly suppressed and supraoptimally facilitated as a function of surround orientation, and these effects diminish as stimulus contrast decreases.

Animals↗

Combined Hebbian development of geniculocortical and lateral connectivity in a model of primary visual cortex.

We present a network model of visual map development in layer 4 of primary visual cortex. Our model comprises excitatory and inhibitory spiking neurons. The input to the network consists of correlated spike trains to mimick the activity of neurons in the lateral geniculate nucleus (LGN). An activity-driven Hebbian learning mechanism governs the development of both the network's lateral connectivity and feedforward projections from LGN to cortex. Plasticity of inhibitory synapses has been included into the model so as to control overall cortical activity. Even without feedforward input, Hebbian modification of the excitatory lateral connections can lead to the development of an intracortical orientation map. We have found that such an intracortical map can guide the development of feedforward connections from LGN to cortical simple cells so that the structure of the final feedforward orientation map is predetermined by the intracortical map. In a scenario in which left- and right-eye geniculocortical inputs develop sequentially one after the other, the resulting maps are therefore very similar, provided the intracortical connectivity remains unaltered. This may explain the outcome of so-called reverse lid-suture experiments, where animals are reared so that both eyes never receive input at the same time, but the orientation maps measured separately for the two eyes are nevertheless nearly identical.

Action Potentials↗

Light and electron microscopic immunocytochemical analysis of the serotonin innervation of the rat visual cortex.

The serotonin afferents of the rat visual cortex were examined immunocytochemically at the light and electron microscopic levels. Immunoreactive fibres were typically thin, tortuous and varicose. Occasionally, some thicker fibres were found. The orientation of labelled axons varied according to laminar position, with fibres running parallel to the pial surface present mainly in layers I and VI, and radially oriented fibres prominent in layers II and III. Branches arising from horizontal or radially oriented fibres were seen to form irregularly shaped loops particularly in layers IV and V. The density of innervation and the prevailing axonal orientation in each cortical layer were similar in both coronal and parasagittal planes. The ultrastructural features of serotonin-labelled axon terminals were examined in single and serial ultrathin sections. While in single sections the majority did not exhibit synaptic specializations, extensive serial section analysis showed that virtually all of these terminals were engaged in junctional complexes. Postsynaptic elements were spines and dendritic shafts, including pyramidal cell apical dendrites, with both symmetrical and asymmetrical membrane specializations. In axospinous synapses, the labelled terminals were usually adjacent to unstained axon terminals contacting the same postsynaptic element.

Animals↗

A theory of the visual motion coding in the primary visual cortex.

This paper demonstrates that much of visual motion coding in the primary visual cortex can be understood from a theory of efficient motion coding in a multiscale representation. The theory predicts that cortical cells can have a spectrum of directional indices, be tuned to different directions of motion, and have spatiotemporally separable or inseparable receptive fields (RF). The predictions also include the following correlations between motion coding and spatial, chromatic, and stereo codings: the preferred speed is greater when the cell receptive field size is larger, the color channel prefers lower speed than the luminance channel, and both the optimal speeds and the preferred directions of motion can be different for inputs from different eyes to the same neuron. These predictions agree with experimental observations. In addition, this theory makes predictions that have not been experimentally investigated systematically and provides a testing ground for an efficient multiscale coding framework. These predictions are as follows: (1) if nearby cortical cells of a given preferred orientation and scale prefer opposite directions of motion and have a quadrature RF phase relationship with each other, then they will have the same directional index, (2) a single neuron can have different optimal motion speeds for opposite motion directions of monocular stimuli, and (3) a neuron's ocular dominance may change with motion direction if the neuron prefers opposite directions for inputs from different eyes.

Color Perception↗

["Timers" and "scanners" among orientation detectors of the visual cortex in the cat].

Acute experiments on immobilized cats were carried out to check the suggestion that a special class of neurons ("timers") which do not change time properties of responses with variations in light bar orientation exists in the visual cortex. Previously neurons of the visual cortex with a dynamic shift of preferable orientation ("scanners") were revealed. Among 76 neurons 27 (36%) showed different properties: stability of latencies in the orientation range, shorter initial and peak latencies, duration and time of the discharge frequency increase, wider orientational tuning and worse relations between magnitudes of responses to preferred and nonpreferred stimuli. They were considered as representatives of the suggested "timers".

Animals↗

Gangliosides fail to enhance behavioral recovery after bilateral ablation of the visual cortex.

Postsurgical injections of GM1 gangliosides (30 mg/kg IP) reduced neither behavioral deficits in rats following bilateral ablation of the visual cortex nor the extent of retrograde degeneration of neurons in the dorsolateral geniculate nucleus that typically accompanies large lesions of the visual cortex. Our findings are in contrast to previous research, in which ganglioside treatments have been shown to enhance the rate of functional recovery after lesions in other parts of the central nervous system. The negative findings in the present experiment may be due to the disruption of normal circadian rhythms caused by occipital cortex injury.

Animals↗

Bilateral projections from the visual cortex to the striatum in the cat.

Direct projections from visual areas 17, 18, 19, and lateral suprasylvian visual area (LS) to the striatum were searched for in 12 adult cats using the autoradiographic technique to detect neuronal pathways. Striatal labels were found only after injections in areas 19 and LS. Projections homolateral to the injection sites were observed from both areas to the head and body of the caudate nucleus and to the putamen. Contralateral projections were found from both areas 19 and LS: however, area 19 did not project to the contralateral putamen. The extent of contralateral projections was smaller and they were confined within the same regions as the homolateral ones. Silver grains were often arranged in cluster-like patches, which were more evident ipsilaterally, in the head of the caudate nucleus and after injections in area LS. The present data support the view of a not strictly topographical segregation of striatal projections from the cat visual cortex.

Animals↗

Hebbian synapses in visual cortex.

We discovered in slices of rat visual cortex that reliable long-term potentiation (LTP) of synaptic responses in layer III could be elicited by theta burst stimulation delivered to a site in the middle of the cortical thickness, corresponding mainly to layer IV. This synaptic plasticity was reflected in the extracellular field potentials and intracellular EPSPs in layer III, but was not observed in the intracellular responses of layer V neurons, suggesting a preferential involvement of synapses on layer III neurons. Tetanus-induced LTP in this preparation was input specific, and was blocked by application of an NMDA receptor antagonist (but not by an antagonist of nitric oxide synthase). In addition, LTP of layer IV-evoked responses could also be produced reliably by pairing low-frequency synaptic stimulation (approximately 100 pulses at 1 Hz) with strong intracellular depolarization of layer III neurons. Thus, LTP in this circuit satisfies the definition of a "Hebbian" modification. Tetanic stimulation of the white matter, in sharp contrast, consistently failed to elicit LTP in layer III unless a GABAA receptor antagonist was applied to the slice. Analysis indicated that the critical difference between layer IV and white matter stimulation was not the magnitude of the responses to single stimuli delivered to the two sites, but that it might lie in the postsynaptic response during high-frequency stimulation. Consistent with this idea, "associative" LTP could be elicited from white matter when converging but independent inputs from the white matter and layer IV simultaneously received tetanic conditioning stimulation. A hypothetical model is presented to account for the differences between layer IV and white matter stimulation. According to this "plasticity gate hypothesis," inhibitory circuitry in layer IV normally acts as a sort of band-pass filter that constrains the types of activity patterns that can gain access to the modifiable synapses in layer III. By stimulating in layer IV, we have bypassed this filter and therefore do not need to block GABAA receptors to achieve the threshold for LTP in layer III.

Action Potentials↗

Cortical cartography revisited: A frequency perspective on the functional architecture of visual cortex.

Viewed in the plane of the cortical surface, the visual cortex is composed of overlapping functional maps that represent stimulus features such as edge orientation, direction of motion, and spatial frequency. Spatial relationships between these maps are thought to ensure that all combinations of stimulus features are represented uniformly across the visual field. Implicit in this view is the assumption that feature combinations are represented in the form of a place code such that a given pattern of activity uniquely signifies a specific combination of stimulus features. Here we review results of experiments that challenge the place code model for the representation of feature combinations. Rather than overlapping maps of stimulus features, we suggest that patterns of activity evoked by complex stimuli are best understood in the context of a single map of spatiotemporal energy.

Animals↗

Critical period plasticity of kitten visual cortex is not associated with enhanced susceptibility to electrical kindling.

The goal of this study was to determine whether the use-dependent malleability of visual cortex functions which is particularly pronounced in 4-week-old kittens correlates with enhanced susceptibility to kindling. For that purpose the effects of high-frequency electrical stimulation were compared in the visual cortex of 4-week-old kittens and of adult cats. The striate cortex of one hemisphere was stimulated with a single train of pulses whose intensity was set just above the threshold for the elicitation of afterdischarges (ADs). In kittens the AD thresholds were consistently higher than in adults and with repeated stimulation, the ADs tended to disorganize, to decrease in amplitude and duration and to become more restricted to the site of stimulation after about 6 stimulations. In the adult, by contrast, the ADs remained well organized and constant in duration throughout 30 stimulations. They showed an increase in amplitude and spike frequency and spread with increasing consistency to the other hemisphere. No electrographic or behavioural signs of epileptic activity developed in kittens, while in adults ADs were on occasion followed by irregular spike activity associated with behavioural states resembling absences. We conclude that the visual cortex possesses powerful mechanisms to prevent the development of supracritical excitatory states, these mechanisms being more effective in the kitten than in the adult.

Age Factors↗

Long-term potentiation of synaptic transmission in kitten visual cortex.

1. Potentiation of synaptic transmission in visual cortex (areas 17 and 18) of kittens was investigated by extracellular recording of field potentials (FPs) and cortical units in cortical slices and whole-animal preparations. Responses to test stimulation (0.05 Hz) of the white matter (WM), lateral geniculate nucleus (LGN), and optic chiasm (OC) were documented before and after conditioning stimulation (2 Hz for 1 h). 2. In slice preparations of area 17, the FPs were always depressed during conditioning stimulation and were usually potentiated immediately after conditioning stimulation. Long-term potentiation (LTP) of FPs developed rapidly during the initial 1-2 h and continued to increase slowly for several hours after conditioning. 3. LTP of FPs was age dependent: LTP occurred most frequently (43/53) at the ages of 21-34 days, less frequently (4/7 and 5/11) at 14-20 and 35-41 days, and never (0/5 and 0/5) at 7-13 and 42-49 days. LTP age relationship determined as a ratio of the amplitudes of FPs after conditioning to that before conditioning was greater at 21-34 days (mean potentiation, 2.4 +/- 0.6) than at 14-20 or 35-41 days (1.7 +/- 0.5). 4. LTP was also documented by the shortening in latencies of orthodromic responses of cortical units sampled from 10 pairs of conditioned and unconditioned control slices. Unit responses were classified into mono- and polysynaptic groups according to the central delay, defined as the time required for their activation after the arrival of afferent impulses. The monosynaptic central delays were 0.22 ms shorter in conditioned (0.60 +/- 0.17 ms, n = 56) than in control slices (0.82 +/- 0.22 ms, n = 57); similarly, polysynaptic central delays were 0.66 ms smaller (1.70 +/- 0.43 ms, n = 51; and 2.36 +/- 0.79 ms, n = 51). Both differences were statistically significant (P less than 0.001). 5. There were laminar differences in LTP of mono- and polysynaptic transmission. LTP of monosynaptic transmission occurred throughout layers II-V (central delays shortened about 0.2 ms), whereas LTP of polysynaptic transmission was greatest in layer II (1.17 ms), moderate in layer III (0.66 ms), and slight in layer IV (0.3 ms). The time course of shortening in orthodromic latency in five polysynaptic units agreed with the time course of LTP of FP. 6. Location of synapses involved in LTP of synaptic transmission was studied by current source-density (CSD) analysis in slice preparations of area 17 during test stimulation of WM. CSD analysis demonstrated two components of current sinks (early and late), probably representing mono- and polysynaptic transmission.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Cerebral vascular malformations adjacent to sensorimotor and visual cortex. Functional magnetic resonance imaging studies before and after therapeutic intervention.

BACKGROUND AND PURPOSE: It is not known how cerebral vascular malformations affect the function of the surrounding brain. Functional magnetic resonance imaging (fMRI) can provide information about normal functional neuroanatomy and its alteration by vascular lesions and therapeutic intervention. METHODS: We performed fMRI studies in 24 patients harboring vascular malformations adjacent to primary somatosensory, motor, and visual cortex. The fMRI studies consisted of the acquisition of an image time series coupled with functional activation of motor, sensory, or visual cortex in both hemispheres. Activated voxels were identified using frequency domain analyses, and their number and anatomic location were compared between the affected and unaffected hemispheres. RESULTS: Every patient capable of performing the desired task showed functional activation. Eight patients without neurological deficits showed a symmetrical pattern of activation between the hemispheres. Each had a vascular malformation located one or more gyri from the functional region imaged. Three patients showed hemispheric symmetry in the location of activated cortex but with a marked asymmetry in the number of activated voxels. Each harbored vascular malformations located within one gyrus of the functional region and showed either subtle or no neurological deficit. Eleven patients showed hemispheric asymmetry in the location of activated cortex. In 6, the anatomic displacement appeared to be due to a mass effect of the lesion. In 5, the activation occurred at a different anatomic locale, and the patients exhibited gross neurological deficit in the respective function. Posttherapeutic changes in functional activation reflected elimination of the mass effect or recovery of clinical function. CONCLUSIONS: Systematic fMRI studies are possible in patients with vascular malformations in brain regions adjacent to primary somatosensory, motor, and visual cortex. Displacement of the activated region and hemispheric asymmetry in the number of activated voxels in the functional regions appear to reflect the anatomic and physiological impact of the vascular malformation. Changes in fMRI findings after intervention reflect the consequences of therapy and parallel clinical recovery.

Adolescent↗

Hypothalamic and basal forebrain afferents to he cat's visual cortex: a study with horseradish peroxidase.

Following injections of horseradish peroxidase into the visual cortex labeled neurons have been found in the nucleus of the diagonal band of Broca in the basal forebrain and in the lateral hypothalamus just rostral to the mammillary bodies. The projection is bilateral, with a strong prevalence of ipsilateral connections. Both the basal forebrain and the lateral hypothalamus project to each of the areas 17, 18 and 19; none of these projections, however, is retinotopically organized. This suggests that the basal forebrain and the lateral hypothalamus are sources of non-specific input to the visual cortex in the cat.

Afferent Pathways↗

Retinotopic maps in human prestriate visual cortex: the demarcation of areas V2 and V3.

We have used PET (positron emission tomography) to chart the mapping of the retina in human occipital visual cortex and hence to locate the secondary and tertiary visual areas, V2 and V3. A group of four non-selected male volunteers was presented with dynamic stimuli that were aligned with either the vertical or the right horizontal meridians (VM or HM) from 0 degree to 29 degrees eccentricity; the vertical stimuli were restricted to either the inferior or the superior hemifields. PET scans were performed using intravenous infusion of H215O and a Siemens-CTI 953B PET scanner with 3D data acquisition. Subjects received 18 scans, divided equally among the right HM, the superior VM, and the inferior VM. Data were analyzed with SPM software. The group average result confirmed our experimental hypothesis that human occipital visual cortex has retinotopic maps similar to those of the macaque monkey. Thus human areas V2 and V3 can be defined on the basis that the border between them is formed by the HM and that the outer border of V3 is demarcated by a second representation of the VM that runs approximately parallel to the primary representation of the VM at the V1/V2 border. Furthermore, as in many mammals, the extrastriate representation of the HM is "split", such that the superior contralateral quadrant is mapped in lower V2 and V3, occupying the ventral surface of human cortex, and the inferior contralateral quadrant is mapped in upper V2 and V3, which extend over the lateral and medial surfaces of each hemisphere. After stereotaxic normalization, the position of V3 defined by retinal topography was found to correspond to that surmised from our previous PET studies employing moving stimuli.

Adult↗

The representation of the horizontal meridian in the primary visual cortex.

The authors report the findings of two patients that confirm the location of the horizontal meridian in the human visual cortex. The first patient had an inferior quadrant defect with a band of horizontal meridian sparing. Magnetic resonance imaging showed a lesion concentrated along the medial striate cortex. The second patient had a homonymous horizontal defect that resulted from removal of an arteriovenous malformation located in the lateral striate cortex. The findings of these two patients demonstrate that the horizontal meridian is represented at the calcarine fissure base in the primary visual cortex.

Adult↗

A historical review of the representation of the visual field in primary visual cortex with special reference to the neural mechanisms underlying macular sparing.

This article comprises a historical review of the literature pertaining to the representation of the visual field in human primary visual cortex. A brief survey of the anatomy of the visual system is followed by a critical evaluation of the key studies that have informed both the issue of the disproportionate representation of central vision within primary visual cortex, and the anatomical basis underlying the phenomena of macular sparing and macular splitting hemianopia.

Hemianopsia↗

High metabolic activity in the visual cortex of early blind human subjects.

Glucose metabolism has been studied in the visual cortex of early blind human subjects. In the forebrain of these subjects, regional glucose utilization was the highest in the striate and prestriate cortical areas. Furthermore, this activity was higher than in blindfolded sighted subjects, whether at rest or during an auditory or tactile task. These observations raise the question of the functionality of the blind's visual cortex.

Blindness↗

Postnatal development of corticocortical efferents from area 17 in the cat's visual cortex.

We are interested in the postnatal development of corticocortical connections in the cat's visual cortex. In this study, we injected the anterograde tracer 3H-proline into visual cortical area 17 of kittens, aged 4-70 d, and adult cats to visualize the distribution of terminals of the association projections to areas 18, 19, 21a, and the lateral suprasylvian visual cortex. The density of anterograde label was quantified using computerized image analysis. There was dense labeling at topographically appropriate locations in area 18 in animals of all ages. In 4- and 8-d-old kittens, other extrastriate areas (19, 21a and the lateral suprasylvian cortex) contained only sparse label, localized in a few solitary axons; these areas were densely labeled in animals aged 12 d or more. In kittens aged 4-20 d there was considerable, widespread label within fibers located in the white matter, and many of these axons lay underneath regions of extrastriate, and also striate, cortex that were almost certainly not destined to be persistently innervated by cells at the injection site. This pattern of extensive white matter label was not seen in animals older than 20 d. In each extrastriate region, from the earliest age at which we identified dense cortical innervation from area 17, the terminals were distributed in clusters. At first these patches were mainly in infragranular layers, but later, during the second and third postnatal weeks, they began to appear in more superficial laminae. By 70 d, an adult-like distribution of terminals was found in each extrastriate area: most fibers appeared to end in layers II and III in areas 18, 19, and 21a and centered on layer IV in the medial bank of the middle suprasylvian sulcus in adult cats. We suggest that the development of ipsilateral association projections from area 17 to extrastriate cortex is a 2-stage process. First, cells at a particular point in area 17 send immature fibers in a nonspecific fashion through white matter towards a very wide area of extrastriate cortex. Second, corticocortical axons penetrate extrastriate cortex mainly in patches at topographically appropriate regions and grow to their targets in a specific fashion.

Aging↗