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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↗

Interhemispheric synchronization of oscillatory neuronal responses in cat visual cortex.

Neurons in area 17 of cat visual cortex display oscillatory responses that can synchronize across spatially separate columns in a stimulus-specific way. Response synchronization has now been shown to occur also between neurons in area 17 of the right and left cerebral hemispheres. This synchronization was abolished by section of the corpus callosum. Thus, the response synchronization is mediated by corticocortical connections. These data are compatible with the hypothesis that temporal synchrony of neuronal discharges serves to bind features within and between the visual hemifields.

Animals↗

Two periods of processing in the (circum)striate visual cortex as revealed by transcranial magnetic stimulation.

To determine the timing of visual processing in the (circum)striate visual cortex, we examined the effect of single pulse transcranial magnetic stimulation over the occipital pole of healthy subjects who were engaged in a forced-choice visual letter identification task. Single letters, subtending a visual angle of 0.35 degrees, were foveally presented for 10 ms and were immediately followed by a mask. We investigated 30 different delays (d), defined as the time between the onset of the visual stimulus and the onset of the magnetic stimulus, from d = -100 ms to d = +190 ms, and 4 different midsagittal coil positions (x), defined as the distance between the lower edge of the coil and the upper edge of the inion, from x = 1 cm to x = 7 cm. Three out of four subjects showed three distinct delay intervals (dips) at which application of TMS resulted in an impairment of the task. The first dip was centred around d = -50 ms and occurred independently of the coil position; the second dip was centred around d = 0 ms and was elicitable only with the two lowest coil positions; the third dip was centred around d = 100 ms and was also elicitable only with the lower coil positions. In the fourth subject, only the first and the third dip were found. We conclude that there are two distinct periods when the activity in the (circum)striate visual cortex is necessary for the identification of visually presented letters.

Electromagnetic Phenomena↗

The rat claustrum: afferent and efferent connections with visual cortex.

We have examined the afferent and efferent projections between the claustrum and visual cortex in the Long-Evans rat using anterograde and retrograde axonal transport techniques. Injections of either wheat germ agglutinin/horseradish peroxidase (WGA/HRP) or Fast Blue were made into each of the main visual regions (17, 18a or 18b) as well as directly into the claustrum. The cortical injections were placed in either the upper, middle or deep layers so as to assist in determining the laminar organization of these connections. Of the 3 visual areas, only area 18b appears to have extensive and reciprocal connections with the claustrum. After a WGA/HRP injection of this area, dense labeled terminals and numerous labeled cells were found intermixed throughout the full extent of the claustrum. The density of this labeled activity was found to vary directly with the amount of the infragranular layers involved by the injections. Injections in the other visual areas did produce labeled cells in the claustrum, but their number was always small or even negligible. There was never any evidence of anterograde labeled terminals in the claustrum from any injection of areas 17 or 18a. Tracer injections directly in the claustrum confirmed and extended these findings by showing that the labeled terminals and/or labeled cells were localized predominantly in layer VI of area 18b of visual cortex. On the basis of these injections, two major conclusions are reached. First, the pattern of connections between the claustrum and visual cortex in the rat differs fundamentally with that found in other species.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Postnatal development of biotinylated dextran amine-labeled corpus callosum axons projecting from the visual and auditory cortices to the visual cortex of the rat.

The distribution and morphology of developing corpus callosum (CC) axons in rat visual cortex was studied by unilateral application of the in vivo anterograde tracer biotinylated dextran amine (BDA) to the visual or auditory cortex of newborns through adults. Changes in the distribution and morphology of CC axons during development were observed. Following BDA placement only in visual cortex, nearly all CC projections were to visual cortex (homotopic CC projections). At postnatal day (PND) 5-8, labeled CC axons were found throughout the contralateral visual cortex, including area 17; these CC axons could be followed from the white matter to layer I. By PND 13, few CC axons were found in medial area 17, indicating the existence of transitory CC axons in area 17 at younger ages. Morphological changes were investigated at the area 17/18a border and showed that CC axon collaterals were not formed until PND 8, and terminal arbors were not visible until PND 13; by PND 17, the adult CC-axon terminal pattern was present. At all ages, only a few heterotopic CC projections from visual to auditory cortex were found in the gray matter, although many labeled CC axons extended laterally into the white matter underlying the auditory cortex. Following BDA placement only in auditory cortex, CC projections to both auditory (homotopic CC projections) and visual (heterotopic CC projections) cortex were observed. At all ages, the homotopic CC projections were present throughout the auditory cortex, but were not distributed homogeneously; densely labeled CC axons showed a distinct columnar organization. The heterotopic CC projections were present in all visual cortical areas, including medial area 17, in significant numbers until PND 24, but were mostly eliminated by PND 28, at which time a labeling pattern similar to the adult was found. Thus, most of the heterotopic CC projections were transitory. The present study confirms the existence of transitory CC axons projecting through all layers of the visual cortex, as revealed by DiI, and extends the DiI results by showing that these transitory CC axons arise from both homotopic and heterotopic origins. Furthermore, different sources of transitory CC axons have different timetables for elimination.

Age Factors↗

Physiological studies in deafferented visual cortex cells of cats following transplantation of fetal xenografts from the rat's cortex.

We have studied the physiological properties of cells (N = 822) in visual cortex area 17 of seven adult cats transplanted with visual cortex xenografts from fetal (E15-E17) rats. The transplants were assumed to induce recovery of adjacent neurons partially deafferented from visual input. The control group (eight cats, 564 cells) had just analogous sectioning in the cortex. The level of activity found, following visual stimulation, in the deafferented cortical region (medially to the graft) was 48.6% compared to the result (34.5%) obtained in the control cats, indicating the preservation of visual responsiveness. Furthermore, no disturbance could be found in the ocular dominance distribution or binocularity (64.4%) of the cells in the grafted region compared to those in the control cats, indicating preservation of the columnar organization. The deafferented cells in the grafted cortex thus demonstrated the absence of adverse immunological reaction there due to the presence of the xenogeneic tissue, indicating that the visual cortex is immunologically privileged.

Afferent Pathways↗

Critical period-dependent reduction of the permissiveness of cat visual cortex tissue for neuronal adhesion and neurite growth.

During postnatal development, the visual cortex undergoes an experience-dependent refinement of its circuitry. This process includes synapse formation, as well as synapse elimination. Both mechanisms appear to be restricted to a limited 'critical period' which lasts for approximately 2 months in cats. We tested whether the termination of the critical period for cortical malleability is paralleled by changes in the growth permissiveness of the tissue. These changes may inhibit progressive reorganization of functional circuitries mediated by axon growth. Embryonic cortical neurons were cultured on unfixed cryostat sections of the visual cortex obtained from cats aged 2-50 weeks. After 2-3 days in vitro the distribution of viable cells and the percentage of neurite-bearing cells were determined and analysed with respect to the developmental age and subdivisions of the underlying tissue substrate. It was shown that cell adhesion and neurite formation are correlated with the developmental age of the substrate tissue and the time period of myelination. While embryonic neurons adhered and survived on grey and white matter tissue from 2- and 4-week-old kittens, there was a significant reduction in cell adhesion on the myelinated white matter regions of the tissue sections of older animals. Quantitative analyses showed that neurite formation by cultured neurons also became successively impaired on grey and white matter areas of tissue substrates, corresponding to the time course of the critical period for cortical malleability. On grey matter tissue this effect was most pronounced between the second and sixth postnatal weeks. The effects were not antagonized by coating the substrate sections with the growth-promoting molecule laminin. It is therefore proposed that neurite growth-inhibiting factors, most probably associated with central nervous system myelin, are gradually expressed postnatally and may contribute to the termination of the critical period in the visual cortex of cats.

Animals↗

Object-based attention in the primary visual cortex of the macaque monkey.

Typical natural visual scenes contain many objects, which need to be segregated from each other and from the background. Present theories subdivide the processes responsible for this segregation into a pre-attentive and attentive system. The pre-attentive system segregates image regions that 'pop out' rapidly and in parallel across the visual field. In the primary visual cortex, responses to pre-attentively selected image regions are enhanced. When objects do not segregate automatically from the rest of the image, the time-consuming attentive system is recruited. Here we investigate whether attentive selection is also associated with a modulation of firing rates in area V1 of the brain in monkeys trained to perform a curve-tracing task. Neuronal responses to the various segments of a target curve were simultaneously enhanced relative to responses evoked by a distractor curve, even if the two curves crossed each other. This indicates that object-based attention is associated with a response enhancement at the earliest level of the visual cortical processing hierarchy.

Animals↗

Functional organization in the visual cortex of the golden hamster.

The visual cortex of the golden hamster was studied by means of multi-unit and single unit recording, which revealed three separate retinotopic maps of the visual field in the posterior cortex. V1, corresponding to cyto-architectonic area 17, has the contralateral temporal field represented medially, the central visual field (extending about 10 deg ipsilateral) represented laterally and the lower field anteriorly. The borders of the map, especially for the upper field, seem to be more restricted than the whole visual field available to the contralateral hemiretina: V1 probably does not represent the extreme periphery of the field. A large fraction of V1 has binocular input, for up to about 50 deg lateral to the vertical midline. There is a retinotopic reversal near the representation of the vertical midline where V1 meets V2 (corresponding to the more lateral "area 18a"). There is another retinotopic reversal at the extremity of the contralateral field representation, where V1 meets Vm (the medial visual area, corresponding to "area 18"). V2 and Vm each contain a reduced mirror image version of the map in V1. Almost all isolated single units in V1 have receptive fields that can be classified as radially symmetrical (60%) or asymmetrical (35%). Symmetrical fields have ON (13%), OFF (4%), ON-OFF (30%) or "SILENT" (12%) central areas when plotted with flashing spots. There are minor but not striking differences between these groups in their field sizes, velocity preferences and so on. They almost invariably prefer moving to stationary stimuli but are not selective for orientation or direction of movement. Asymmetrical fields are of four types, three of which (type 1, 11%; type 2, 17%; and type 3, 2%) are orientation selective and resemble simple, complex and hypercomplex cells in the cat cortex. Some of these have direction as well as orientation preference. Axial movement detectors (5%) have a selectivity for one axis of motion, and thus prefer one orientation of edge, but respond equally well to movement of a spot. Vertical and horizontal orientation preferences, especially the latter, are much the most common. There is some evidence of clustering of cells according to receptive field type and, possibly, preferred orientation. Asymmetrical cells are, relatively somewhat rarer in the deeper cortical layers. Within the binocular segment, fully 89% of cells are binocularly driven and the receptive fields are similar in the two eyes. Receptive fields tend to increase in size away from the area centralis representation and, in a complementary fashion, the magnification factor decreases from up to 0.1 mm/deg at the area centralis representation to about 0.02 mm/deg for the peripheral field.

Animals↗

Orientation formed by a spot's trajectory: a two-dimensional population approach in primary visual cortex.

There exist a large number of visual illusions indicating that perception differs from pure representation of physical input. For example, a spot of light can be characterized by its position, but it does not contribute any information about orientation. However, when moved fast enough, a continuous streak along its trajectory is perceived that helps to determine the orientation of the movement path. The question arises whether the processing of the trajectory and its orientation are simultaneously represented in the primary visual cortex. Here I show that decoding neural population activity within a two-dimensional parameter space represents both (1) physical input given by the actual position of the moving spot and (2) orientation. This latter parameter has no physical counterpart in the stimulus but must be actively formed by spatiotemporal integration of the spot's trajectory.

Action Potentials↗

Disparity tuning and binocularity of single neurons in cat visual cortex.

Activity from single neurons in the visual cortex of anaesthetized and paralyzed cats were recorded in response to monocular and binocular stimuli of different retinal disparities. Three different types of disparity sensitive neurons were found, characterized by (1) a response maximum near zero disparity, (2) a response minimum near zero disparity, and (3) an asymmetric disparity sensitivity. Usually, neurons of the first type receive about the same monocular input from the two eyes; the other disparity sensitive neurons respond better to monocular stimulation of one eye as compared to the other. The results closely resemble those obtained recently in the rhesus monkey.

Animals↗