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Distribution and morphology of functionally identified neurons in the visual cortex of the rat.

The distribution and morphology of functionally identified neurons were examined in the visual cortex of Long Evans pigmented rats. The results, based on qualitative and quantitative analysis of single cell spike activity, have shown that neurons in the rat visual cortex have well-defined receptive field properties and are similar to those reported for animals with more highly developed visual systems. Unlike the cat and monkey, the distribution of receptive field types appeared even throughout the visual cortex. Exception was provided by layer IV which, similar to the more 'visual' animals, contained the largest percentage of simple cells. Horseradish peroxidase injected into single, physiologically identified neurons allowed for detailed morphological characterization of functional cell types. Of the cells successfully filled with horseradish peroxidase, complex cells were pyramidal in morphology and located in layers II through VI. Simple cells were both pyramidal and non-pyramidal in appearance and were located in layers II + III and IV. Finally, hypercomplex cells were pyramidal in appearance and their perikarya were situated in layers II + III and V.

Animals↗

Amphetamine-induced recovery of visual cliff performance after bilateral visual cortex ablation in cats: measurements of depth perception thresholds.

After bilateral visual cortex ablation, cats exhibit a loss of depth perception as measured on a visual cliff, which recovers following administration of d-amphetamine. In this Study, 3 amphetamine-treated cats with visual cortex ablations showed a rapid and enduring recovery, with 2 of these animals obtaining levels of performance seen only with binocular vision, suggesting a restoration of binocular depth perception. Cats with asymmetrical lesions showed only a transient improvement during amphetamine treatment, and some animals not displaying autonomic signs of amphetamine intoxication did not improve. Saline-treated cats showed no signs of improvement, and the effect of amphetamine was blocked by the catecholaminergic antagonist haloperidol. These results indicate that amphetamine can induce an enduring recovery from a behavioral deficit after brain injury, which if left untreated would not spontaneously recover.

Amphetamine↗

Computational design and nonlinear dynamics of a recurrent network model of the primary visual cortex.

Recurrent interactions in the primary visual cortex make its output a complex nonlinear transform of its input. This transform serves preattentive visual segmentation, that is, autonomously processing visual inputs to give outputs that selectively emphasize certain features for segmentation. An analytical understanding of the nonlinear dynamics of the recurrent neural circuit is essential to harness its computational power. We derive requirements on the neural architecture, components, and connection weights of a biologically plausible model of the cortex such that region segmentation, figure-ground segregation, and contour enhancement can be achieved simultaneously. In addition, we analyze the conditions governing neural oscillations, illusory contours, and the absence of visual hallucinations. Many of our analytical techniques can be applied to other recurrent networks with translation-invariant neural and connection structures.

Animals↗

Early extrastriate activity without primary visual cortex in humans.

Damage to the primary visual cortex (V1) destroys the major source of anatomical input to extrastriate cortical areas (V2, V3, V4 and V5) and produces cortical blindness--an absence of any sensation of light and colour--in the visual field contralateral to the side of the lesion. Neuroimaging studies, nevertheless, have recently demonstrated dorsal and ventral extrastriate activation for stationary stimuli presented to the blind visual field in the absence of V1 activity in human subjects. To clarify the moment in time that visual information reaches extrastriate areas, by means of event-related potentials (ERPs) we tracked the temporal course of responses to complex visual stimuli (faces) presented in the blind field of a hemianopic patient. Stimulation of the normal visual field elicited a positive occipital deflection (P1) at 140 ms. A P1 response was also observed with stimulation of the blind field, although slightly delayed (20 ms) and reduced. Its topography and timing demonstrate that early neural activity for stationary stimuli takes place within extrastriate regions despite V1 denervation.

Adult↗

Differential effects of quinolinic acid lesions on muscarinic acetylcholine receptors in cat visual cortex during postnatal development.

Quinolinic acid (QA) lesions of neurons in cat visual cortex were combined with conventional in vitro autoradiographic methods in order to define the cellular locus of the muscarinic acetylcholine receptor (mAChR). Animals of various postnatal ages had QA unilaterally injected into the visual cortex. Four to fourteen days later they were sacrificed and processed for electron microscopy (EM) or in vitro autoradiography. QA lesions at the various postnatal ages were found to eliminate intrinsic cortical neurons and their processes while leaving intact glia, fibers of passage and axon terminals from outside the lesion zone. Autoradiograms of visual cortex labelled with [3H]QNB (which labels M1 and M2 subtypes) showed an age-dependent loss of binding sites, with the greatest decreases occurring after 65 days postnatal. Examined separately, only the M1 mAChRs labelled with [3H]pirenzepine exhibited these age-dependent alterations. The results indicate a differential distribution of the M1 mAChRs during postnatal development. The loss of receptors late in postnatal life following QA suggests a dominantly neuronal locus; the relatively small loss early in postnatal life suggests a locus on other cellular elements.

Aging↗

[Differences in the properties of 2 groups of orientation detectors of the visual cortex in the cat].

Functional properties of 149 neurons in field 17 of the visual cortex were studied in immobilized cats. They were divided into two groups. In 37% of neurons the initial and peak latencies of responses changed not more than by 10 ms in the orientation range. They were named "timer"-neurons. 63% of neurons were called "scanners". "Timers" reliably differed from "scanners" in shorter latencies, time of the discharge frequency increase, duration of the response and higher discharge frequency in all stimuli orientations. Scanners had a higher orientation selectivity, they considerably more often scanned the range of orientations in time. Neurons of the two groups had contraphase dynamics of the width of tuning orientation during response while distribution of their preferential orientations supplemented each other. In orientation columns of the visual cortex, both timers and scanners were most often found with a predominance of the latter. The columns consisting of only timers or only scanners were more seldom. The role of differences in properties of two neuronal groups in the cat visual cortex for orientation detection is discussed.

Animals↗

Functional organization of the cat visual cortex in relation to the representation of a uniform surface.

Neuronal activity in the early visual cortex has been extensively studied from the standpoint of contour representation. On the other hand, representation of the interior of a surface surrounded by a contour is much less well understood. Several studies have identified neurons activated by a uniform surface covering their receptive fields, but their distribution within the cortex is not yet known. The aim of the present study was to obtain a better understanding of the distribution of such neurons in the visual cortex. Using optical imaging of intrinsic signals, we found that there are a group of surface-responsive regions located in area 18, along the area 17/18 border, that tend to overlap the singular points of the orientation-preference map. Extracellular recordings confirmed that neurons responsive to uniform plane stimuli are accumulated in these regions. Such neurons also existed outside the surface-responsive regions around the singular points. These results suggest that there exists a functional organization related to the representation of a uniform surface in the early visual cortex.

Animals↗

How does the cerebral cortex work? Learning, attention, and grouping by the laminar circuits of visual cortex.

The organization of neocortex into layers is one of its most salient anatomical features. These layers include circuits that form functional columns in cortical maps. A major unsolved problem concerns how bottom-up, top-down, and horizontal interactions are organized within cortical layers to generate adaptive behaviors. This article models how these interactions help visual cortex to realize: (i) the binding process whereby cortex groups distributed data into coherent object representations; (ii) the attentional process whereby cortex selectively processes important events; and (iii) the developmental and learning processes whereby cortex shapes its circuits to match environmental constraints. New computational ideas about feedback systems suggest how neocortex develops and learns in a stable way, and why top-down attention requires converging bottom-up inputs to fully activate cortical cells, whereas perceptual groupings do not.

Attention↗

Experience-dependent plasticity of mouse visual cortex in the absence of the neuronal activity-dependent marker egr1/zif268.

Neuronal activity elicits a rapid increase in the expression of several immediate early genes (IEGs). To clarify a role for IEG response in activity-dependent development, we examined the contribution of the egr1/zif268 gene during visual cortical processing and plasticity in mice. We first analyzed the expression of egr1 mRNA in wild-type (WT) mice using Northern blot hybridization. In the visual cortex, expression of egr1 mRNA increased dramatically after eye opening, systemic injection of kainate, or 30 min of photostimulation after a brief (5 d) period of dark adaptation. Thus, the expression of egr1 is regulated by synaptic activity in the mouse visual cortex, as it is in other species (e.g., monkeys, cats, and rats). To evaluate whether this transcription factor is directly involved in activity-dependent plasticity, mice lacking Egr1 were deprived of the use of one eye during the developmental critical period [postnatal day 24 (P24)-P34]. Extracellular in vivo single-unit recordings from the binocular zone of the visual cortex revealed that visual responses developed normally in egr1 knock-out (KO) mice. Moreover, a similarly significant shift of responsiveness in favor of the open eye was produced in both KO and WT mice by either brief (4 d) or long-term (>2 weeks) occlusion of one eye. There was no apparent compensation among egr2, egr3, or c-fos mRNA and protein expression in the visual cortex of egr1 KO mice. Taken together, these results indicate that egr1 is a useful marker of sensory input in mice but is not intrinsically necessary for the experience-dependent plasticity of the visual cortex. Our findings underscore a mechanistic distinction between sensory plasticity and long-lasting forms of synaptic potentiation in the hippocampus, for which egr1/zif268 was recently found to be essential.

Aging↗

Muscarinic acetylcholine receptor subtypes in rat visual cortex--a comparative study using quantitative receptor autoradiography and in situ hybridization.

The laminar pattern of M1- and M2-muscarinic acetylcholine receptors (mAChR) in rat visual cortex has been compared with the distribution of the corresponding m1, m2, m3 and m4 receptor genes using both quantitative receptor autoradiography and in situ hybridization histochemistry. The laminar distribution of 3H-pirenzepine binding to M1-mAChRs in rat visual cortex shows a bimodal pattern with higher binding levels in upper layer III and deeper layer VI. In contrast, highest binding of 3H-AF-DX384 to M2-mAChRs was observed in upper layer IV (100%) and upper layer VI (about 80% of highest binding). The m1 receptor mRNA is almost homogeneously distributed throughout the visual cortex, whereas the m2mAChr mRNA predominates in layer IV with lower levels in layers I and V. The highest amounts of m3mAChR mRNA in rat visual cortex were observed in layer II, while the distribution of m4mAChR transcripts shows a bimodal pattern with peaks in layers III and upper layer VI. The distinct laminar pattern of mRNA muscarinic receptor subtypes in rat visual cortex suggest specific roles of the muscarinic receptor in visual function.

Animals↗

Evidence for a laminar organization of basal forebrain afferents to the visual cortex.

The present study shows that restriction of HRP injections to layer I within the visual cortex results in negligible retrograde labeling within the nuclei of the basal forebrain. In contrast, when the injections of either HRP or WGA-HRP are restricted to the granular and infragranular layers of visual cortex, extensive retrograde labeling occurs within the basal forebrain. Based upon these findings, we argue that the projection from the basal nucleus terminates preferentially within the deep layers of the visual cortex, and thus contributes minimally to the supragranular layers, including layer I.

Acetylcholinesterase↗

Image features selected by neurons of the cat primary visual cortex.

The sensitivity of neurons in field 17 of the visual cortex in cats to cross-shaped, Y-shaped, and star-shaped figures flashing in the receptive field was studied. About 40% of the neurons studied (114 of 289) were found to generate large responses (with an average response factor of 3.06 +/- 0.32) to one of the figures flashing in the center of the receptive field, as compared with the responses produced to a single bar in the optimal orientation. Most of these neurons (72%) were selectively sensitive to the shape and orientation of figures; the remainder demonstrated some degree of tuning invariance to these properties. The latent periods of responses to figures were usually shorter than those of responses to bars. Tuning parameters for bars and figures were generally related: neurons with acute orientational tuning to a bar were usually highly selective to both the configuration and the orientation people figures. Separate or combined stimulation with crosses in the center and near periphery of the receptive fields demonstrated summation, antagonism, or the lack of any interaction between these zones in producing sensitivity to crosses. Local blockade of intracortical GABAergic inhibition by microiontophoretic application of bicuculline showed that in one third of the neurons studied, sensitivity to figures was generated or enhanced by inhibition in normal conditions, while one third of cells showed suppression by inhibition, and sensitivity in the remainder was independent of inhibition. These data show that reconsideration of existing concepts of the role of field 17 in selecting only first-order shape features of images (i.e., the orientations of single lines) is needed, since almost half the neurons in the cat primary visual cortex can efficiently detect second-order features (angles and line intersections).

Action Potentials↗

Functional MRI of visual cortex in sedated 18 month-old infants with or without periventricular leukomalacia.

Functional MRI (fMRI) of the visual cortex was evaluated in 42 sedated 18-month-old infants (mean corrected age; 31 males, 11 females) with or without periventricular leukomalacia (PVL). Data from 14 infants could not be evaluated because of movement artefacts. Ten of the remaining 28 infants showed no significant fMRI response upon visual stimulation. In 18 infants, a significant signal change upon stimulation was found in the visual cortex: in 17 a signal decrease and in one a signal increase. Functional changes were located mainly in the anterior part of the visual cortex. Seven of the 28 infants had normal MRI and 21 showed variable occipital PVL. An fMRI response was equally frequent in infants without PVL (4 of 7 infants) and with PVL (14 of 21 infants). In conclusion, fMRI was shown to be feasible in sedated infants. No correlation was found between functional activation and the presence or absence of occipital PVL. Type of fMRI response (signal decrease) and localization (anterior part of the visual cortex) are different from those seen in adults, probably reflecting a combination of sedation effects and immaturity of the visual system. At present, fMRI is a highly promising research tool; its clinical relevance still has to be established.

Case-Control Studies↗

Involvement of nerve growth factor in visual cortex plasticity.

The physiological role of nerve growth factor (NGF), the prototype member of the neurotrophin family, has been widely studied. NGF has been shown to promote survival, sprouting and differentiation of sympathetic ganglion cells and sensory neurons in the peripheral nervous system; it has also been shown to support survival and regeneration of cholinergic neurons in the central nervous system. Recent evidence indicates that NGF is also involved in the neuronal plasticity of the visual cortex. Exogenous supplies of NGF have been shown to interfere with normal processes underlying activity- and age-dependent synaptic modifications in both developing and adult visual cortex. In parallel to these physiological effects, numerous neuronal markers in the visual cortex have been found to be influenced by NGF. Several proposals have been introduced to explain the physiological role of NGF in visual cortex plasticity. Although the mechanisms underlying NGF effects in the visual cortex are still under active investigation, current evidence implies that NGF, and perhaps other neurotrophins as well, may be useful for preventing or correcting inappropriate or anomalous connections in the visual cortex, and thus for treating visual dysfunctions such as amblyopia and strabismus.

Animals↗

Neural field model of receptive field restructuring in primary visual cortex.

Receptive fields (RF) in the visual cortex can change their size depending on the state of the individual. This reflects a changing visual resolution according to different demands on information processing during drowsiness. So far, however, the possible mechanisms that underlie these size changes have not been tested rigorously. Only qualitatively has it been suggested that state-dependent lateral geniculate nucleus (LGN) firing patterns (burst versus tonic firing) are mainly responsible for the observed cortical receptive field restructuring. Here, we employ a neural field approach to describe the changes of cortical RF properties analytically. Expressions to describe the spatiotemporal receptive fields are given for pure feedforward networks. The model predicts that visual latencies increase nonlinearly with the distance of the stimulus location from the RF center. RF restructuring effects are faithfully reproduced. Despite the changing RF sizes, the model demonstrates that the width of the spatial membrane potential profile (as measured by the variance sigma of a gaussian) remains constant in cortex. In contrast, it is shown for recurrent networks that both the RF width and the width of the membrane potential profile generically depend on time and can even increase if lateral cortical excitatory connections extend further than fibers from LGN to cortex. In order to differentiate between a feedforward and a recurrent mechanism causing the experimental RF changes, we fitted the data to the analytically derived point-spread functions. Results of the fits provide estimates for model parameters consistent with the literature data and support the hypothesis that the observed RF sharpening is indeed mainly driven by input from LGN, not by recurrent intracortical connections.

Electroencephalography↗

Low-threshold Ca2+ channels mediate induction of long-term potentiation in kitten visual cortex.

1. The induction mechanism of long-term potentiation (LTP) in developing visual cortex was studied by recording intracellular responses from layer III-IV cells in slice preparations of kitten visual cortex at 30-40 days after birth. 2. Strong stimulation of white matter produced a late depolarizing response after an orthodromic action potential. This depolarizing response was abolished by membrane depolarization or hyperpolarization caused by current injection through the recording electrode. In addition, this response was reduced by bath application of a low concentration (100 microM) of Ni2+ without any changes in the rising slope of the excitatory postsynaptic potential (EPSP) or orthodromic action potential. This suggests that this response is mediated by low-threshold Ca2+ channels (LTCs). 3. The involvement of LTCs in the induction of LTP was tested. White matter was stimulated at 2 Hz for 15 min as a conditioning stimulus to induce LTP, and the resultant changes in EPSPs were tested by low-frequency (0.1 Hz) stimulation of white matter. Conditioning stimulation produced a large N-methyl-D-aspartate (NMDA) receptor-mediated depolarizing response in these cells, which obscured the presence of the late depoliarzation. Therefore the test was conducted in a solution containing an NMDA antagonist 2-amino-5-phosphonovalerate (APV). 4. Weak conditioning stimulation, which evoked no LTC responses, never induced LTP; whereas strong conditioning stimulation, which evoked LTC responses, always induced LTP. Strong conditioning stimulation failed to induce LTP when LTC responses were prevented either by membrane depolarization or hyperpolarization or by a bath application of 100 microM Ni2+. 5. In a solution without APV, the application of Ni2+ also prevented the induction of LTP. 6. When cells were impaled by an electrode containing a Ca2+ chelator 1,2-bis-(o-aminophenoxy)ethane-N,N,N',N'-tetraacetic acid (BAPTA), LTP was never induced, even though LTC responses were evoked by conditioning stimulation. These results indicate that Ca2+ influx into postsynaptic cells through LTCs induces the LTP. 7. The responses mediated by LTCs, which were evoked by the injection of current pulses into the cells, were maximum at the critical period of visual cortical plasticity, suggesting that LTCs in postsynaptic cells regulate the plastic changes in developing visual cortex.

2-Amino-5-phosphonovalerate↗

Test-retest analysis with functional MR of the activated area in the human visual cortex.

PURPOSE: To investigate the intersubject and intrasubject variability of the activated area in the visual cortex with functional MR imaging. METHODS: Double-section gradient-echo MR images were acquired at 1.5 T in 28 healthy volunteers using the fast low-angle shot (FLASH) technique. Visual stimulation was obtained with light-emitting diode (LED) goggles. Eighteen volunteers were studied twice. The size of the activated areas in the visual cortex and the increase in signal were measured. A reproducibility ratio for size (R[size]) and for location (R[overlap]) was calculated on a scale of 0.0 to 1.0. RESULTS: Activation was seen in 89% of the subjects. The size of the activated area was widely variable among subjects: mean, 460 +/- 284 mm2; range, 0 to 1029 mm2 in the first study in all volunteers. Signal increases ranged from 3.2% to 10.9%, with a mean of 6.6 +/- 1.7%. The mean values of intrasubject variability testing were R(size) = 0.83 +/- 0.16 and R(overlap) = 0.31 +/- 0.11. CONCLUSION: Functional MR imaging with the FLASH technique is useful in identifying certain cortical areas that have quite variable locations among subjects. This study provides reference data for the intrasubject and intersubject variability of the activation pattern of the visual cortex.

Adult↗

Human EEG responses to 1-100 Hz flicker: resonance phenomena in visual cortex and their potential correlation to cognitive phenomena.

The individual properties of visual objects, like form or color, are represented in different areas in our visual cortex. In order to perceive one coherent object, its features have to be bound together. This was found to be achieved in cat and monkey brains by temporal correlation of the firing rates of neurons which code the same object. This firing rate is predominantly observed in the gamma frequency range (approx. 30-80 Hz, mainly around 40 Hz). In addition, it has been shown in humans that stimuli which flicker at gamma frequencies are processed faster by our brains than when they flicker at different frequencies. These effects could be due to neural oscillators, which preferably oscillate at certain frequencies, so-called resonance frequencies. It is also known that neurons in visual cortex respond to flickering stimuli at the frequency of the flickering light. If neural oscillators exist with resonance frequencies, they should respond more strongly to stimulation with their resonance frequency. We performed an experiment, where ten human subjects were presented flickering light at frequencies from 1 to 100 Hz in 1-Hz steps. The event-related potentials exhibited steady-state oscillations at all frequencies up to at least 90 Hz. Interestingly, the steady-state potentials exhibited clear resonance phenomena around 10, 20, 40 and 80 Hz. This could be a potential neural basis for gamma oscillations in binding experiments. The pattern of results resembles that of multiunit activity and local field potentials in cat visual cortex.

Adult↗