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Early versus late visual cortex lesions: effects on receptive fields in cat superior colliculus.

Cats that sustain lesions of the visual cortex early in life appear to perform certain visual discrimination tasks better than those operated as adults. This study sought to determine whether this recovery is visual capacities was accompanied by reorganization of single cell responses at the level of the superior colliculus. Areas 17 and 18 were ablated in adult cats and in kittens at various times during the neonatal period. Responses of units in superior colliculus ipsilateral to the lesion were recorded following a prolonged recovery period. Following cortical lesions, collicular units rarely exhibited direction selectivity, binocularity was reduced in the majority of animals, and the ocular dominance distribution was biased toward the contralateral eye. The reduction of direction selectivity and binocularity were unrelated to the animal's age at operation.

Age Factors↗

Serotonin facilitates synaptic plasticity in kitten visual cortex: an in vitro study.

We have addressed the role of serotonin-2C (5-HT2C) receptors in the development and maintenance of synaptic plasticity in the kitten visual cortex. In visual cortical slices, taken from 40- to 80-day-old kittens, bath application of serotonin markedly facilitated the induction of both long-term depression (LTD) and long-term potentiation (LTP). Field potential responses to white matter stimulation were recorded from layer IV after a regime of low frequency stimulation (LFS; 1 Hz, 15 min), which reliably induced LTP or LTD in younger kittens (less than 30 days of age). At 40-80 days, this protocol almost never induced LTD or LTP in layer IV. However, in 50% of the visual cortical slices studied in 40-80-day-old kittens, LTD or LTP was induced, if serotonin (1 or 10 microM) was co-applied with LFS. No such serotonin facilitation of long-term plasticity was ever detected in > 120-day-old animals, indicating that serotonin facilitates synaptic plasticity within a defined period of visual cortical development. Serotonergic 5-HT2C receptors are likely to contribute to the synaptic plasticity observed in layer IV, since mesulergine, an antagonist of the 5-HT2C receptor, completely blocked synaptic modifications induced by the combination of low frequency stimulation and serotonin application.

Aging↗

Optical imaging of intrinsic signals as a tool to visualize the functional architecture of adult and developing visual cortex.

One of the most common principles of cortical organization is that neurons with similar response properties are clustered together in space. Thereby the environment is represented in an orderly fashion on the cortical surface in a so-called "cortical map". In primary visual cortex, for instance, neurons with similar orientation preferences are grouped together, forming the orientation preference map. Optical imaging of intrinsic signals allows to investigate the organization of such maps in vivo. Neuronal activity was measured utilizing the fact that the transition from oxy-hemoglobin to hemoglobin in active brain areas can be detected optically by recording changes in light reflectance with a high resolution CCD-camera. When using this technique to look at the exact patterning of orientation preference maps in cat visual cortex a novel principle for the organization of cortical maps was observed: orientation was not organized in parallel bands as had previously been thought but iso-orientation domains were organized radially; orientations from 0 to 180 degrees were laid out in a pinwheel-like fashion around singularities which we termed "orientation-centers". After observing pinwheel patterns in orientation preference maps in adult cat visual cortex it was also investigated how these meticulously arranged maps develop in the cortex of young kittens. Performing chronical recordings in kittens from the age of postnatal day 17 on we were able to observe how orientation maps form already during the third week of life and--under normal conditions--remain largely unchanged thereafter.(ABSTRACT TRUNCATED AT 250 WORDS)

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Visual cortex ablations do not prevent extinction of fear-potentiated startle using a visual conditioned stimulus.

Following observations in the literature that sensory cortex ablations prevent extinction of conditioned fear, the present experiments tested the generality of this finding by examining whether visual cortex ablations would prevent extinction of conditioned fear as assessed by fear-potentiated startle using a visual conditioned stimulus. Consistent with previous reports, visual cortex ablations did not prevent the acquisition or expression of fear-potentiated startle to a visual conditioned stimulus. More importantly, visual cortex ablations did not prevent extinction of fear-potentiated startle to a visual conditioned stimulus, nor did they reverse preoperatively established extinction, indicating that sensory cortex is not required for extinction of conditioned fear in all situations.

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[Testing the scanning effect of a range of orientations by visual cortex neurons of the cat].

The preservation of the effect of dynamic scanning of a part of the orientation diapason during development of neuron responses in the primary visual cortex of cat was examined after equalization of their latencies and after estimation of only highly significant fragments of their reactions. It was found that this effect was preserved in 13 neurons out of 17 examined ones: it remained invariable in 7 units of this group and was modified due to a shift of the scanning beginning along the diapason of orientations, to an increase of scanning diapason or to a change of scanning direction. The stabilization of the orientational tuning occurred only in 4 cells after equalization of latencies. The obtained results confirm that the dynamic changes of orientation tuning in the majority of visual cortex neurons are connected with reorganization of the time pattern of the reaction, as suggested by the hypothesis of the spatial-temporal orientation coding in the visual system.

Animals↗

Acute effects of alcohol on photic evoked potentials of albino rats: visual cortex and superior colliculus.

Photic evoked potentials were recorded from the primary visual cortex (VC) and superior colliculus (SC) of chronically implanted rats. Animals were given intraperitoneal injections of saline, 1.5 and 2.5 g ethyl alcohol/kg body weight on separate days. Evoked potentials were recorded at 5, 20, 40 and 60 min following injection. The amplitudes of all of the VC components except P2 (latency of 52 msec) were depressed to some extent by both doses of alcohol. In contrast, the amplitude of component P2 was increased by both alcohol doses. In the SC, the peak amplitudes of two individual components of the early positive complex were diminished by both doses of alcohol, as was a later negative component. A series of late oscillatory potentials recorded from the SC were minimally depressed by the 1.5 g/kg dose of alcohol, but showed a more prolonged depressant effect at the higher dose. Both doses of alcohol produced reliable increases in peak latency for the primary components in the VC and SC.

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Spatial periodicities of periodic complex cells in the visual cortex cluster at one-half octave intervals.

Within individual penetrations in the visual cortex, spatial periodicities of periodic complex cells differ by either one-half or one octave. When data are pooled from neurons subserving the central visual area in many cats, the results indicate that spatial periodicities cluster at one-half octave intervals over a 2 1/2-octave range (0.95 to 5.4 cyc/deg). Thus a relatively small number of such channels spaced at regular intervals along a logarithmic scale within each orientation column may suffice for this stage of spatial processing.

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Orientation selectivity in visual cortex by fluctuation-controlled criticality.

Within a large-scale neuronal network model of macaque primary visual cortex, we examined how intrinsic dynamic fluctuations in synaptic currents modify the effect of strong recurrent excitation on orientation selectivity. Previously, we showed that, using a strong network inhibition countered by feedforward and recurrent excitation, the cortical model reproduced many observed properties of simple and complex cells. However, that network's complex cells were poorly selective for orientation, and increasing cortical self-excitation led to network instabilities and unrealistically high firing rates. Here, we show that a sparsity of connections in the network produces large, intrinsic fluctuations in the cortico-cortical conductances that can stabilize the network and that there is a critical level of fluctuations (controllable by sparsity) that allows strong cortical gain and the emergence of orientation-selective complex cells. The resultant sparse network also shows near contrast invariance in its selectivity and, in agreement with recent experiments, has extracellular tuning properties that are similar in pinwheel center and iso-orientation regions, whereas intracellular conductances show positional dependencies. Varying the strength of synaptic fluctuations by adjusting the sparsity of network connectivity, we identified a transition between the dynamics of bistability and without bistability. In a network with strong recurrent excitation, this transition is characterized by a near hysteretic behavior and a rapid rise of network firing rates as the synaptic drive or stimulus input is increased. We discuss the connection between this transition and orientation selectivity in our model of primary visual cortex.

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Intrinsic connections in cat visual cortex: a combined anterograde and retrograde tracing study.

Area 18 of cat visual cortex was examined for intrinsic axons following small, columnar injections of an anterograde tracer, Phaseolus vulgaris leucoagglutinin (PHA-L). Locally projecting axons radiated from the injection site and branched to form 10-15 discrete, approximately circular patches 500-750 microns in diameter consisting of many bouton-studded terminal arborizations. Labeled fibers and boutons ramified densely in layers I, II/II, V, and VI, and were noticeably less dense in layer IV. Afferent and efferent pathways originating from the same cortical columns were studied by injecting a mixture of PHA-L and wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP). Between 10 and 15 patches of cells retrogradely labeled by WGA-HRP surrounded each injection site. Within a patch, labeled cells were found in all layers and included both pyramidal and non-pyramidal cells. The distribution of PHA-L labeling was similar to that obtained when PHA-L was injected alone. Most often, the labeled patches resulting from injections of such mixtures contained both anterograde and retrograde labeling. However, patches consisting of retrograde labeling alone and of anterograde labeling alone were also observed, indicating that the local connections linking neighboring cortical columns were not always reciprocal.

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Gain control from beyond the classical receptive field in primate primary visual cortex.

Gain control is a salient feature of information processing throughout the visual system. Heeger (1991, 1992) described a mechanism that could underpin gain control in primary visual cortex (V1). According to this model, a neuron's response is normalized by dividing its output by the sum of a population of neurons, which are selective for orientations covering a broad range. Gain control in this scheme is manifested as a change in the semisaturation constant (contrast gain) of a V1 neuron. Here we examine how flanking and annular gratings of the same or orthogonal orientation to that preferred by a neuron presented beyond the receptive field modulate gain in V1 neurons in anesthetized marmosets (Callithrix jacchus). To characterize how gain was modulated by surround stimuli, the Michaelis-Menten equation was fitted to response versus contrast functions obtained under each stimulus condition. The modulation of gain by surround stimuli was modelled best as a divisive reduction in response gain. Response gain varied with the orientation of surround stimuli, but was reduced most when the orientation of a large annular grating beyond the classical receptive field matched the preferred orientation of neurons. The strength of surround suppression did not vary significantly with retinal eccentricity or laminar distribution. In the marmoset, as in macaques (Angelucci et al., 2002a,b), gain control over the sort of distances reported here (up to 10 deg) may be mediated by feedback from extrastriate areas.

Animals↗

Feature-based attention increases the selectivity of population responses in primate visual cortex.

BACKGROUND: Attending to the spatial location or to nonspatial features of visual stimuli can modulate neuronal responses in primate visual cortex. The modulation by spatial attention changes the gain of sensory neurons and strengthens the representation of attended locations without changing neuronal selectivities such as directionality, i.e., the ratio of responses to preferred and anti-preferred directions of motion. Whether feature-based attention acts in a similar manner is unknown. RESULTS: To clarify this issue, we recorded the responses of 135 direction-selective neurons in the middle temporal area (MT) of two macaques to an unattended moving random dot pattern (the distractor) positioned inside a neuron's receptive field while the animals attended to a second moving pattern positioned in the opposite hemifield. Responses to different directions of the distractor were modulated by the same factor (approximately 12%) as long as the attended direction remained unchanged. On the other hand, systematically changing the attended direction from a neuron's preferred to its anti-preferred direction caused a systematic change of the attentional modulation from an enhancement to a suppression, increasing directionality by about 20%. CONCLUSIONS: The results show that (1) feature-based attention exerts a multiplicative modulation upon neuronal responses and that the strength of this modulation depends on the similarity between the attended feature and the cell's preferred feature, in line with the feature-similarity gain model, and (2) at the level of the neuronal population, feature-based attention increases the selectivity for attended features by increasing the responses of neurons preferring this feature value while decreasing responses of neurons tuned to the opposite feature value.

Animals↗

Neuroplasticity after unilateral visual cortex damage in the newborn cat.

Anatomical, electrophysiological, and behavioral studies implicate extrastriate cortex as a major contributor to the sparing of visually guided behaviors following lesions of primary visual cortex incurred early in life. Here we report considerable sparing of the ability to detect and localize stimuli in the hemifield contralateral to unilateral early lesions of all contiguous visually-responsive primary and extrastriate cortical regions (occipital, visuoparietal, and visuotemporal cortices). In the adult cat this same lesion induces a dense blindness and cats are unable to orient to any visual stimulus introduced into the contralesional hemifield. In the absence of cortical circuits, the neural sparing identified following the neonatal lesion is based on the superior colliculus and it occurs despite massive retrograde transynaptic degeneration of large numbers of retinal ganglion cells.

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Linearity of summation of synaptic potentials underlying direction selectivity in simple cells of the cat visual cortex.

Intracellular recordings from simple cells of the cat visual cortex were used to test linear models for the generation of selectivity for the direction of visual motion. Direction selectivity has been thought to arise in part from nonlinear processes, as suggested by previous experiments that were based on extracellular recordings of action potentials. In intracellular recordings, however, the fluctuations in membrane potential evoked by moving stimuli were accurately predicted by the linear summation of responses to stationary stimuli. Nonlinear mechanisms were not required.

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Effects of neurotrophins on synaptic protein expression in the visual cortex of dark-reared rats.

Total lack of visual experience [dark rearing (DR)] is known to prolong the critical period and delay development of sensory functions in mammalian visual cortex. Recent results show that neurotrophins (NTs) counteract the effects of DR on functional properties of visual cortical cells and exert a strong control on critical period duration. NTs are known to modulate the development and synaptic efficacy of neurotransmitter systems that are affected by DR. However, it is still unknown whether the actions of NTs in dark-reared animals involve interaction with neurotransmitter systems. We have studied the effects of DR on the expression of key molecules in the glutamatergic and GABAergic systems in control and NT-treated animals. We have found that DR reduced the expression of the NMDA receptor 2A subunit and its associated protein PSD-95 (postsynaptic density-95), of GRIP (AMPA glutamate receptor interacting protein), and of the biosynthetic enzyme GAD (glutamic acid decarboxylase). Returning dark-reared animals to light for 2 hr restored normal expression of the above-mentioned proteins almost completely. NT treatment specifically counteracts DR effects; NGF acts primarily on the NMDA system, whereas BDNF acts primarily on the GABAergic system. Finally, the action of NT4 seems to involve both excitatory and inhibitory systems. These data demonstrate that different NTs counteract DR effects by modulating the expression of key molecules of the excitatory and inhibitory neurotransmitter systems.

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The visual cortex of the agouti (Dasyprocta aguti): architectonic subdivisions.

1. We have studied the cytoarchitecture and myeloarchitecture of the agouti's cortical surface that can be activated by visual stimulation. Five architectonic subdivisions that correspond to distinctive visuotopic representations were characterized. 2. The largest portion of the visual cortex is occupied by area 17 which is situated lateral to the cingulate cortex, medial to area 18, posterior to the parietal cortex, and anterior to the agranular retrosplenial cortex. Additionally, four architectonic subdivisions in the extrastriate visual cortex were distinguished, i.e., from medial to lateral: area 18, area 19, anterior lateral area, and temporal posterior area. 3. Along the border of the extrastriate cortex a ring of nonvisual cortical fields was encountered encompassing parietal (somatic sensorial) cortex, temporal anterior and temporal intermediate (auditory) areas, a band of pre-rhinal cortex, and agranular retrosplenial cortex.

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A GAP-43-like protein in cat visual cortex.

We have purified a protein that changes in relative concentration during the development of the kitten visual cortex. It resembles GAP-43 (a neuronal protein that is expressed at elevated levels during periods of development and regenerative axon growth) in the following respects: (1) it is an acidic protein (pI = 4.7) whose electrophoretic mobility on SDS-PAGE is similar to, but lower than rat GAP-43, suggesting that the cat protein is larger; (2) its electrophoretic mobility varies with the acrylamide concentration in a manner that is characteristic of GAP-43; (3) its concentration in kitten forebrain is elevated during early postnatal development; (4) the sequence of ten consecutive amino acids from a chemically generated fragment matches the expected sequence from GAP-43; and (5) its amino-acid content also matches GAP-43. We conclude that our purified protein is cat GAP-43. Immunoblots with an antibody prepared against rat GAP-43 suggested that the concentration of GAP-43 in the visual cortex declines with age.

Acrylamide↗

The formation of cooperative cell assemblies in the visual cortex.

During a critical period of postnatal development of the mammalian visual cortex, synaptic connections are susceptible to use-dependent modifications. Synaptic connections strengthen if pre- and postsynaptic elements are active simultaneously and postsynaptic depolarization is sufficient to allow for the activation of N-methyl-D-aspartate (NMDA)-receptor-gated conductances. By contrast, synaptic gain decreases if postsynaptic activation exceeds a critical threshold and presynaptic afferents are not capable of activating NMDA-receptor-dependent conductances. These processes lead to selective stabilization of connections between neuronal elements which often exhibit correlated activity and thus modify connectivity according to functional criteria. It is suggested that such experience-dependent selection of circuits serves different purposes at different levels of visual processing. At the input stage to the striate cortex it contributes to optimize the match between the representations of the two eyes. At a later stage of processing it participates in the development of selective connections between cortical columns and thereby serves to establish neuronal representations for frequently occurring constellations of features. Use-dependent changes of synaptic gain can also be induced in the mature visual cortex. These modifications follow the same rules as those occurring during early development and appear to depend on similar molecular mechanisms. However, in the adult the changes of synaptic gain do not seem to be followed by major rearrangements of connectivity. This suggests developmental alterations in mechanisms responsible for growth, removal and stabilization of synaptic connections. Actually, many of the cellular mechanisms thought to be involved in use-dependent synaptic plasticity change during development but it is still unclear which of them are responsible for the definitive stabilization of functionally confirmed pathways.

Aging↗

Optical mapping of electrical activity in rat somatosensory and visual cortex.

We have investigated the use of optical methods for monitoring neuron activity in mammalian cortex. The cortex was stained with a voltage-sensitive dye and fluorescence was simultaneously measured from 124 areas using a photodiode array. Optical signals were detected in rat somatosensory cortex in response to small whisker movements and in visual cortex in response to light flashes to the eye. Relatively large signals were obtained during focal interictal epileptiform discharges induced by bicuculline. The measuring system had a time resolution of milliseconds and a spatial resolution of a few hundred micrometers. Simultaneous, multi-site optical recordings of activity may provide a new and potentially powerful method for studying function and dysfunction in mammalian cortex.

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