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PubMed2001

AMPA receptor properties at the synapse between retinal afferents and thalamocortical cells in the dorsal lateral geniculate nucleus of the rat.

The thalamocortical (TC) cells in dorsal lateral geniculate nucleus transfer signals from retinal afferents to the primary visual cortex. The excitatory retinal input to the TC cells is mediated by ionotropic receptors of the N-methyl-D-aspartate (NMDA) and non-NMDA type. In the present study the excitatory postsynaptic current (EPSC) mediated by non-NMDA receptors in this synapse was characterised by means of voltage-clamp recordings from TC neurons in rat thalamic slices. The specific alpha-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid receptor antagonist GYKI-53655 fully blocked the non-NMDA mediated EPSC, evoked by optic tract stimulation. The EPSC peak amplitudes were linearly related to the command voltage, suggesting that the receptor complex includes the GluR2 subunit. The EPSC amplitude and decay time increased during application of the desensitisation blocker, cyclothiazide, showing that the EPSC was partly controlled by fast desensitisation.

2-Amino-5-phosphonovalerate↗
PubMed2003

Occ1 mRNA expression reveals a characteristic feature in the hippocampal CA2 field of adult macaques.

The gene occ1 is preferentially expressed in the primary visual cortex in an activity-dependent and developmentally regulated manner. In this report, we show the characteristic distribution of occ1 transcripts in the CA2 subfield of the hippocampal formation in adult monkeys. occ1 mRNA signals were observed selectively in the pyramidal cell layer of CA2. In addition to these signals, a relatively sparse distribution of occ1 was found in the stratum oriens and, occasionally, in the outermost regions of the pyramidal cell layers of both CA1 and CA2. A few labeled cells were detected in CA3. The elevated expression of occ1 in the CA2 subfield provides a new approach for investigating the function of this subregion, whose role has still not been well clarified.

Animals↗
PubMed1996

The functional basis of ocular dominance: functional MRI (fMRI) findings.

Changes in cortical metabolism and cerebral perfusion may be recorded non-invasively with functional magnetic resonance imaging (fMRI). In pilot experiments, using fMRI with photic stimulation, we found differences between activated areas when the left or the right eye was stimulated separately. In this study we investigated whether this could be explained by ocular dominance. We studied 26 healthy volunteers (mean age 23.3 +/- 3.5 years). Ocular dominance was determined by means of the near-far alignment test. fMRI-measurements consisted of a double-slice gradient echo sequence. Slices were acquired placed parallel on either side of the calcarine fissure. Visual stimulation was done with goggles with two LED matrices (red light, 8 Hz); each in front of one eye. In each subject, the left and right eye were stimulated separately and together, in a randomly alternating order. Twenty-two subjects showed activation, of whom eight subjects had a dominant left eye and 14 a dominant right eye. In general the size of the activated area was bigger upon stimulation of the dominant eye. The difference with the area upon stimulation of the non-dominant eye was statistically significant in the right eye dominant group. These results indicate that the dominant eye actually activates a larger area of the primary visual cortex than the non-dominant eye. This provides for the first time a functional basis for the concept of ocular dominance.

Adult↗
PubMed2002

Spatio-temporal plasticity of cortical receptive fields in response to repetitive visual stimulation in the adult cat.

Many psychophysical experiments on perceptual learning in humans show increases of performance that are most probably based on functions of early visual cortical areas. Long-term plasticity of the primary visual cortex has so far been shown in vivo with the use of visual stimuli paired with electrical or pharmacological stimulation at the cellular level. Here, we report that plasticity in the adult visual cortex can be achieved by repetitive visual stimulation. First, spatial receptive field profiles of single units (n=38) in area 17 or 18 of the anesthetized cat were determined with optimally oriented flashing light bars. Then a conditioning protocol was applied to induce associative synaptic plasticity. The receptive field center and an unresponsive region just outside the excitatory receptive field were synchronously stimulated ('costimulation', repetition rate 1 Hz; for 10-75 min). After costimulation the receptive field and its adjacent regions were mapped again. We observed specific increases of the receptive field size, changes of the receptive field subfield structure as well as shifts in response latency. In 37% of the cells the receptive field size increased specifically towards the stimulated side but not towards the non-stimulated opposite side of the receptive field. In addition, changes in the relative strength and size of the on and off subfield regions were observed. These specific alterations were dependent on the level of neuronal activity during costimulation. During recovery, the new responses dropped down to 120% of the preconditioning value on average within 103 min; however, the decay times significantly depended on the response magnitude after costimulation. In the temporal domain, the latency of new responses appeared to be strongly influenced by the latency of the response during costimulation.Twenty-nine percent of the units displayed no receptive field enlargement, most likely because the activity during costimulation was significantly lower than in the cases with enlarged receptive fields. An unspecific receptive field enlargement towards both the stimulated and non-stimulated side was observed in 34% of the tested cells. In contrast to the cells with specifically enlarged receptive fields, the unspecific increase of receptive field size was always accompanied by a strong increase of the general activity level. We conclude that the receptive field changes presumably took place by strengthening of synaptic inputs at the recorded cells in a Hebbian way as previously shown in the visual cortex in vitro and in vivo. The observed receptive field changes may be related to preattentive perceptual learning and could represent a basis of the 'filling in' of cortical scotomas obtained with specific training procedures in human patients suffering from visual cortex lesions.

Animals↗
PubMed2003

Regional brain cytochrome oxidase activity in beta-amyloid precursor protein transgenic mice with the Swedish mutation.

Cytochrome oxidase activity was examined in a transgenic mouse model of Alzheimer's disease with overexpression of the 751 amino acid isoform of beta-amyloid precursor protein with the Swedish mutation under control of the murine thy-1 promoter. The neuritic plaques, abundantly localized in the hippocampus and anterior neocortical areas, showed a core devoid of enzymatic activity surrounded by higher cytochrome oxidase activity at the sites of the dystrophic neurites and activated glial cells. Quantitative measures, taken only in the healthy-appearing regional areas without neuritic plaques, were higher in numerous limbic and non-limbic regions of transgenic mice in comparison with controls. Enzymatic activity was higher in the dentate gyrus and CA2-CA3 region of the hippocampus, the anterior cingulate and primary visual cortex, two olfactory structures, the ventral part of the neostriatum, the parafascicularis nucleus of the thalamus, and the subthalamic nucleus. Brainstem regions anatomically related with altered forebrain regions were more heavily labeled as well, including the substantia nigra, the periaqueductal gray, the superior colliculus, the medial raphe, the locus coeruleus and the adjacent parabrachial nucleus, as well as the pontine nuclei, red nucleus, and trigeminal motor nucleus. Functional brain organization is discussed in the context of Alzheimer's disease. Although hypometabolism is generally observed in this pathology, the increased cytochrome oxidase activity obtained in these transgenic mice can be the result of a functional compensation on the surviving neurons, or of an early mitochondrial alteration related to increased oxidative damage.

Alzheimer Disease↗
PubMed1998

The significance of precisely replicating patterns in mammalian CNS spike trains.

Neuronal spike trains from both single and multi-unit recordings often contain patterns such as doublets and triplets of spikes that precisely replicate themselves at a later time. The presence of such precisely replicating patterns can still be detected when the tolerance on interval replication is shortened to a fraction of a millisecond. In this context we examine here data taken from various parts of the central nervous systems of anesthetized rats, cats and monkeys. The relative abundance of replicating triplets varies from centre to centre, and is nearly always significantly greater than obtained in Monte-Carlo simulations of either a Poisson-like process or a renewal process having the same interspike interval distribution as the neuronal data. However, a remarkable exception is found in the activity of retinal ganglion cells. Significant deviations were found in the primary visual cortex and, even more so, in the lateral geniculate body and the mitral cells of the olfactory bulb. Using a fixed tolerance for the replication of intervals (0.5 ms) it is usually observed that replicating patterns are produced in excess (with respect to renewal process models) mostly in low firing rate episodes (< or = 100 Hz). However, using a tolerance that varies in direct proportion to the mean interval (i.e. as the reciprocal of the firing rate), one generally observes that replicating triplets occur with higher than expected frequency in comparable proportions at all firing rates. This observation suggests the existence of a scale invariance principle in these phenomena with respect to certain neuronal codes. In order to decrease the influence of the estimated neuronal firing rate on the results of the comparisons, we computed also the ratio NT2/ND3, of the number of replicating triplets to the number of doublets replicating three times [Lestienne R. (1994) Proc. Soc. Neurosci. 20, 22; Lestienne R. (1996) Biol. Cybern. 74, 55-61], using both a fixed or a variable tolerance. In spike trains obeying a Poisson process, NT2/ND3 ratios should be nearly independent of the frequency, especially when using a variable tolerance. These studies supported previous results: significant deviations from the models are found in all the spike trains examined, except in the case of retinal ganglion cells, and the most significant deviations are found in recordings from the lateral geniculate nucleus and the mitral cells of the olfactory bulb. Removing spikes that belong to bursts having large "Poisson surprise" values [Legéndy C. R. and Salcman M. (1985) J. Neurophysiol. 53, 926-939] (except the very first spike of the burst) significantly decreases NT2/ND3 ratios in the record from the lateral geniculate nucleus, suggesting that in this case bursty episodes greatly contribute to the production of replicating patterns, but such a removal does not affect results from the piriform record. Finally, in both the lateral geniculate nucleus and in the mitral cells of the olfactory bulb records, perturbing the timing of spikes by applying to interspike intervals small jitters of uniform probability density with amplitude up to 3 ms, very significantly decrease NT2/ND3 ratios in these centres, but does not change much the NT2/ND3 ratios in other neuronal recordings. Implications of these findings for a possible role of precisely replicating patterns in temporal coding of neuronal information is discussed, as well as possible mechanisms for their production.

Animals↗
PubMed1998

Spatial and temporal matching of receptive field properties of binocular cells in area 19 of the cat.

The spatial and temporal properties of single neurons were investigated in area 19 of the cat. We evaluated the matching of binocular receptive field properties with regard to the respective strength of the ipsilateral and contralateral inputs. Results indicate that most cells in area 19 are well tuned to spatial and temporal frequencies and exhibit relatively low contrast threshold (mean=6.8%) when assessed using optimal parameters and tested through the dominant eye. Spatial resolution (mean=0.75 c/degree), optimal spatial frequencies (mean=0.16 c/degree) were relatively low and spatial bandwidths (mean=2.1 octaves) were broader as compared to those of cells in area 17 but comparable to those of cells in other extrastriate areas. On the other hand temporal resolution (mean=10.7 Hz), optimal temporal frequency (mean=4.5 Hz) and temporal bandwidths (mean=2.9 octaves) were higher and broader than in primary visual cortex. A significant relationship exists between most of the cell's properties assessed through either eye. For some parameters, such as spatial and temporal resolution, ocular dominance was shown to be significantly related to the extent of matching between the two eyes. For these parameters, binocular cells that exhibited a balanced ocular dominance were generally well matched with regard to the receptive field properties of each eye whereas the largest mismatches were found in cells that were more strongly dominated by one eye. These results suggest that visual input contributes to the activation of cells in area 19 in a redundant manner, possibly attesting to the multiplicity of parallel pathways to this area in the cat.

Animals↗
PubMed2001

Influence of baseline hematocrit and hemodilution on BOLD fMRI activation.

Current understanding of blood oxygenation level dependent (BOLD) fMRI physiology predicts a close relationship between BOLD signal and blood hematocrit level. However, neither this relationship nor its effect on BOLD percent activation (BPA) has been empirically examined in man. To that end, BPA in primary visual cortex in response to photic stimulation was determined in a group of 24 normal subjects. A positive linear relationship between BPA and hematocrit was seen, particularly in men. To evaluate the effect of change in hematocrit on BPA, 9 men were studied before and following isotonic saline hemodilution, resulting in an average 6% reduction in hematocrit and an 8-31% reduction in BPA. No significant change in the number of activated pixels was seen. A model of predicted BPA as a function of hematocrit and vessel size was developed, and results from this model closely mirrored the empiric data. These results suggest that hematocrit significantly influences the magnitude of BPA and that such baseline factors should be accounted for when comparing BOLD data across groups of subjects, particularly in the many instances in which hematocrit may vary systematically. Such instances include several disease states as well as studies involving sex differences, drug administration, stress and other factors. Finally, the robust agreement between predicted and empiric data serves to validate a semiquantitative approach to the analysis of BOLD fMRI data.

Brain↗
PubMed2000

Stimulus frequency affects c-fos expression in the rat visual system.

We have characterised the c-fos expression patterns in various centers of the visual pathway of adult rats monocularly stimulated either by continuous or flickering light at different frequencies. Results show different immunocytochemical patterns in all centers studied, the geniculate lateral complex (LGC), superior colliculus (SC) and primary visual cortex (Oc1), depending on the physical characteristics of the stimulus (blinking frequency and light wavelength). After stimulation of the left eye, the ipsilateral pathway presents a substantial density of immunoresponsive cells, which is greater than expected with respect to the number of fibers that project ipsilaterally from the retina to the LGC and the superficial layers of the SC. A surprisingly high positive immunoresponsiveness is obtained in all cases with coherent light stimulation in the red spectrum (634 nm).

Animals↗
PubMed1998

A neural network model for the development of simple and complex cell receptive fields within cortical maps of orientation and ocular dominance.

Prenatal development of the primary visual cortex leads to simple cells with spatially distinct and oriented ON and OFF subregions. These simple cells are organized into spatial maps of orientation and ocular dominance that exhibit singularities, fractures, and linear zones. On a finer spatial scale, simple cells occur that are sensitive to similar orientations but opposite contrast polarities, and exhibit both even-symmetric and odd-symmetric receptive fields. Pooling of outputs from oppositely polarized simple cells leads to complex cells that respond to both contrast polarities. A neural network model is described which simulates how simple and complex cells self-organize starting from unsegregated and unoriented geniculocortical inputs during prenatal development. Neighboring simple cells that are sensitive to opposite contrast polarities develop from a combination of spatially short-range inhibition and high-gain recurrent habituative excitation between cells that obey membrane equations. Habituation, or depression, of synapses controls reset of cell activations both through enhanced ON responses and OFF antagonistic rebounds. Orientation and ocular dominance maps form when high-gain medium-range recurrent excitation and long-range inhibition interact with the short-range mechanisms. The resulting structure clarifies how simple and complex cells contribute to perceptual processes such as texture segregation and perceptual grouping.

Journal Article↗
PubMed1999

Self-organization of shift-invariant receptive fields.

This paper proposes a new learning rule by which cells with shift-invariant receptive fields are self-organized. With this learning rule, cells similar to simple and complex cells in the primary visual cortex are generated in a network. To demonstrate the new learning rule, we simulate a three-layered network that consists of an input layer (or the retina), a layer of S-cells (or simple cells), and a layer of C-cells (or complex cells). During the learning, straight lines of various orientations sweep across the input layer. Here both S- and C-cells are created through competition. Although S-cells compete depending on their instantaneous outputs, C-cells compete depending on the traces (or temporal averages) of their outputs. For the self-organization of S-cells, only winner S-cells increase their input connections in a similar way to that for the neocognitron. In other words, the winner S-cells have LTP (long term potentiation) in their input connections. For the self-organization of C-cells, however, loser C-cells decrease their input connections (LTD=long term depression), while winners increase their input connections (LTP). Here both S- and C-cells are accompanied by inhibitory cells. Modification of inhibitory connections together with excitatory connections is important for creation of C-cells as well as S-cells.

Journal Article↗
PubMed2000

Low response variability in simultaneously recorded retinal, thalamic, and cortical neurons.

The response of a cortical cell to a repeated stimulus can be highly variable from one trial to the next. Much lower variability has been reported of retinal cells. We recorded visual responses simultaneously from three successive stages of the cat visual system: retinal ganglion cells (RGCs), thalamic (LGN) relay cells, and simple cells in layer 4 of primary visual cortex. Spike count variability was lower than that of a Poisson process at all three stages but increased at each stage. Absolute and relative refractory periods largely accounted for the reliability at all three stages. Our results show that cortical responses can be more reliable than previously thought. The differences in reliability in retina, LGN, and cortex can be explained by (1) decreasing firing rates and (2) decreasing absolute and relative refractory periods.

Action Potentials↗
PubMed2001

Stability of cortical responses and the statistics of natural scenes.

The primary visual cortex (V1) of higher mammals contains maps of stimulus features; how these maps influence vision remains unknown. We have examined the functional significance of an asymmetry in the orientation map in cat V1, i.e., the fact that a larger area of V1 is preferentially activated by vertical and horizontal contours than by contours at oblique orientations. Despite the fact that neurons tuned to cardinal and oblique orientations have indistinguishable tuning characteristics, cardinal neurons remain more stable in their response properties after selective perturbation induced by adaptation. Similarly, human observers report different adaptation-induced changes in orientation tuning between cardinal and oblique axes. We suggest that the larger cortical area devoted to cardinal orientations imposes stability on the processing of cardinal contours during visual perception, by retaining invariant cortical responses along cardinal axes.

Adaptation, Physiological↗
PubMed2002

Opponent inhibition: a developmental model of layer 4 of the neocortical circuit.

We model the development of the functional circuit of layer 4 (the input-recipient layer) of cat primary visual cortex. The observed thalamocortical and intracortical circuitry codevelop under Hebb-like synaptic plasticity. Hebbian development yields opponent inhibition: inhibition evoked by stimuli anticorrelated with those that excite a cell. Strong opponent inhibition enables recognition of stimulus orientation in a manner invariant to stimulus contrast. These principles may apply to cortex more generally: Hebb-like plasticity can guide layer 4 of any piece of cortex to create opposition between anticorrelated stimulus pairs, and this enables recognition of specific stimulus patterns in a manner invariant to stimulus magnitude. Properties that are invariant across a cortical column are predicted to be those shared by opponent stimulus pairs; this contrasts with the common idea that a column represents cells with similar response properties.

Action Potentials↗
PubMed2002

Rapid ocular dominance plasticity requires cortical but not geniculate protein synthesis.

Synaptic plasticity is a multistep process in which rapid, early phases eventually give way to slower, more enduring stages. Diverse forms of synaptic change share a common requirement for protein synthesis in the late stages of plasticity, which are often associated with structural rearrangements. Ocular dominance plasticity in the primary visual cortex (V1) is a long-lasting form of activity-dependent plasticity comprised of well-defined physiological and anatomical stages. The molecular events underlying these stages remain poorly understood. Using the protein synthesis inhibitor cycloheximide, we investigated a role for protein synthesis in ocular dominance plasticity. Suppression of cortical, but not geniculate, protein synthesis impaired rapid ocular dominance plasticity, while leaving neuronal responsiveness intact. These findings suggest that structural changes underlying ocular dominance plasticity occur rapidly following monocular occlusion, and cortical changes guide subsequent alterations in thalamocortical afferents.

Animals↗
PubMed2002

S cone contributions to the magnocellular visual pathway in macaque monkey.

The magnocellular visual pathway is believed to receive input from long (L) and middle (M), but not short (S), wavelength-sensitive cones. Recording from neurons in magnocellular layers of lateral geniculate nucleus (LGN) in macaque monkeys, we found that magnocellular neurons were unequivocally responsive to S cone-isolating stimuli. A quantitative analysis suggests that S cones provided about 10% of the input to these cells, on average, while L:M ratios were far more variable. S cone signals influenced responses with the same sign as L and M cone inputs (i.e., no color opponency). Magnocellular afferent recordings following inactivation of primary visual cortex demonstrated that S cone signals were feedforward in nature and did not arise from cortical feedback to LGN

Action Potentials↗
PubMed2002

Autophosphorylation of alphaCaMKII is required for ocular dominance plasticity.

Experience is a powerful sculptor of developing neural connections. In the primary visual cortex (V1), cortical connections are particularly susceptible to the effects of sensory manipulation during a postnatal critical period. At the molecular level, this activity-dependent plasticity requires the transformation of synaptic depolarization into changes in synaptic weight. The molecule alpha calcium-calmodulin kinase type II (alphaCaMKII) is known to play a central role in this transformation. Importantly, alphaCaMKII function is modulated by autophosphorylation, which promotes Ca(2+)-independent kinase activity. Here we show that mice possessing a mutant form of alphaCaMKII that is unable to autophosphorylate show impairments in ocular dominance plasticity. These results confirm the importance of alphaCaMKII in visual cortical plasticity and suggest that synaptic changes induced by monocular deprivation are stored specifically in glutamatergic synapses made onto excitatory neurons.

Aging↗
PubMed2002

Synaptic integration by V1 neurons depends on location within the orientation map.

Neurons in the primary visual cortex (V1) are organized into an orientation map consisting of orientation domains arranged radially around "pinwheel centers" at which the representations of all orientations converge. We have combined optical imaging of intrinsic signals with intracellular recordings to estimate the subthreshold inputs and spike outputs of neurons located near pinwheel centers or in orientation domains. We find that neurons near pinwheel centers have subthreshold responses to all stimulus orientations but spike responses to only a narrow range of orientations. Across the map, the selectivity of inputs covaries with the selectivity of orientations in the local cortical network, while the selectivity of spike outputs does not. Thus, the input-output transformation performed by V1 neurons is powerfully influenced by the local structure of the orientation map.

Action Potentials↗