Development of spatial and temporal selectivity in the suprasylvian visual cortex of the cat

We have studied the development of the spatial and temporal properties of neurons in the medial bank of the suprasylvian visual cortex (PMLS) in kittens aged between 9 d and 8 weeks. Quantitative measurements were made of the responses to drifting high-contrast gratings of optimum orientation and direction of motion, but varying in spatial and temporal frequency. The spatial resolution ("acuity") of cells increased rapidly and was fully mature (over 2 cycles/deg for the best cells) at 3 weeks of age. The optimum spatial frequency also tended to improve and reached adult values (around 0.5 cycles/deg for the best cells) at about the end of the third week. In younger kittens, the spatial resolution of neurons was not obviously correlated with the eccentricity of their receptive fields, but in older animals acuity was clearly elevated for receptive fields in the central visual field. The proportion of "low-pass" cells (showing no obvious attenuation of response for gratings of low spatial frequency) decreased with age and simultaneously there was a slight increase in the mean spatial bandwidth of "bandpass" cells. Responses to drifting sinusoidal gratings were generally dominated by an unmodulated elevation of discharge at all ages. In tests with stationary, contrast-modulated gratings presented at different spatial positions, cells in the youngest kittens behaved nonlinearly and showed mainly an unmodulated increase in discharge, whereas in older kittens, as in adult cats, most neurons responded to contrast-modulated gratings with a small, phase-dependent response at the temporal frequency of modulation and a larger component at twice the fundamental frequency. None of the cells recorded at any age had a true "null position." As in adult PMLS, the widths of receptive fields in kittens were, on average, about twice the size of the preferred spatial period (4 times the preferred bar width). At all ages, therefore, neurons in PMLS resembled striate complex cells with respect to the nonlinearity of their responses and the spatial structure of their receptive fields. The preferred temporal frequency and high-temporal-frequency cutoff also improved, on average, during the first 3 weeks of life, and the range of temporal frequencies over which cells responded continued to increase until at least 8 weeks. Although the low-spatial-frequency inhibition that creates spatial bandpass characteristics probably depends on cortical mechanisms, the postnatal development of both temporal and spatial resolution might well be limited by maturation at the level of the retina.     

Development of neuronal responses in cat posteromedial lateral suprasylvian visual cortex

Exchange assays have often been used to quantitate steroid receptors when endogenous ligands are present; however, there are no reports of their successful application to activated glucocorticoid-Type II receptor complexes. In addition to investigating the reasons for this failure, the present study also examined the effects of progesterone on glucocorticoid dissociation from, and reassociation with unactivated and activated Type II receptors. Molybdate-stabilized brain cytosol from adrenal-ovariectomized mice was incubated with [3H]dexamethasone ( +/- [1H]DEX) for 40 h at 0 degree C. Afterwards free steroid was removed on Sephadex G-25 columns in the presence (unactivated receptors) or absence (activated receptors) of molybdate. Activation, as measured by DNA-cellulose binding, was achieved by incubating molybdate-free cytosol at 22 degrees C for 20 min followed by G-25 filtration in the presence of molybdate. The rates of dissociation and reassociation were then measured by incubating cytosol with [1H]triamcinolone acetonide (TA) or [3H]TA ( +/- [1H]TA) at 12 degrees C. An exchange assay was also employed in which cytosol was incubated first with [1H]DEX for 40 h at 0 degree C followed by bound-free steroid separations and 12 degrees C incubations with [3H]TA ( +/- [1H]TA). Both approaches revealed that even though activation reduced the rate of DEX dissociation from Type II receptors by 40%, it eliminated the ability of the newly unoccupied receptors to rebind glucocorticoid. Adding [1H]progesterone to occupied receptor preparations increased dissociation rate constants by nearly 3-fold, for both unactivated and activated Type II receptors. Since [1H]TA failed to prevent this effect, progesterone appears to act at an allosteric site(s) which cannot be occupied by glucocorticoids. Exchange assays revealed that progesterone-facilitated dissociation increased the rate of glucocorticoid rebinding to unactivated, but not activated Type II receptors. These results suggest that spontaneous and progesterone-facilitated termination of glucocorticoid genomic actions could be mediated by steroid dissociation since unoccupied activated Type II receptors do not rebind agonist steroid.     

Neuronal connections underlying orientation selectivity in cat visual cortex

How do neurons of the visual cortex acquire their acute sensitivity to the orientation of a visual stimulus? The question has preoccupied those who study the cortex since Hubel and Wiesel¹ first described orientation selectivity over twenty-five years ago. At the time, they proposed an elegant and enduring model for the origin of orientation selectivity. Fig. 1A, which is adapted from their original paper and which contains the essence of their model, is by now familiar to most students of the visual system and to many others besides. Yet the model, and the central question that it addresses, is still the subject of intense debate. Competing models have arisen in the intervening years, along with diverse experiments that bear on them.     

Orientation selectivity of visually sensitive neurons of the lateral suprasylvian region of the cat cerebral cortex

Orientation selectivity of single units was found in a lateral suprasylvian area of cat, which has not been described previously. Changes of orientational tuning and preferred orientation in time were revealed by the "time slices" method.     

Spontaneous and induced recovery of visual hemineglect during cooling inactivation of middle suprasylvian cortex in the cat

The purpose of the present study was to observe recovery of orienting deficits during unilateral and bilateral cooling inactivation of the middle suprasylvian (MS) cortex. Unilateral MS cooling resulted in a contralateral hemineglect (no orienting responses towards stimuli in the contracooled hemifield) that spontaneously recovered within less than 40 min of continued cooling. Moreover, the brain changes responsible for this recovery were partially retained and cumulated over successive cooling days. Bilateral MS cooling indicated that recovery of neglect can be induced by a restoration of the imbalance between activity levels of structures in both hemispheres and revealed an interaction between spontaneous and induced recovery of neglect. These results show that considerable changes may occur in the brain during cooling for rather short periods. Possible explanations for these changes are discussed.     

Synaptic organization of cortico-cortical connections from the primary visual cortex to the posteromedial lateral suprasylvian visual area in the cat

The synaptic organization of the projection from the cat striate visual cortex to the posteromedial lateral suprasylvian cortical area (PMLS) was examined. The anterograde tracer Phaseolus vulgaris leucoagglutinin (PHA-L) was iontophorectically delivered into area 17, and anterogradely labeled fibers were revealed in PMLS by means of an immunocytochemical detection method. Most axons and presumptive terminal swellings were found in layers III and IV. The neuronal elements (n = 190) that were postsynaptic to anterogradely labeled boutons were quantitatively analyzed. All anterogradely labeled cortico-cortical boutons (n = 182) established type 1 synapses. The results show that 83% of the postsynaptic targets were dendritic spines, probably belonging to pyramidal cells. Dendritic shafts constituted 17% of the targets. The dendritic shafts postsynaptic to cortico-cortical boutons were studied for the presence of gamma-aminobutyric acid (GABA) with a postembedding immunogold method. Most dendritic shafts (85%) that were tested were found to be GABA-positive, demonstrating that they originate from local inhibitory neurons. Taking into account that most postsynaptic targets were spines and extending the results of the immunocytochemical testing to the total population of postsynaptic dendrites, it was calculated that at least 14% of targets originated from GABA-positive cells. Thus cortico-cortical axons establish direct monosynpatic connections mainly with pyramidal and to a lesser extent with GABAergic nonpyramidal neurons in area PMLS, providing both feedforward excitation and feedforward inhibition to a visual associational area known to be involved in the processing of motion information. The results are consistent with previously demonstrated deficits in physiological properties of neurons in PMLS following removal of cortico-cortical afferents.     

Orientation selectivity in cat primary visual cortex: local and global measurement

In this study,we investigated orientation selectivity in cat primary visual cortex(V1) and its relationship with various parameters.We found a strong correlation between circular variance(CV) and orthogonal-topreferred response ratio(O/P ratio),and a moderate correlation between tuning width and O/P ratio.Moreover,the suppression far from the peak that accounted for the lower CV in cat V1 cells also contributed to the narrowing of the tuning width of cells.We also studied the dependence of orientation selectivity on the modulation ratio for each cell,which is consistent with robust entrainment of the neuronal response to the phase of the drifting grating stimulus.In conclusion,the CV(global measure) and tuning width(local measure) are significantly correlated with the modulation ratio.     

Comparison of the selectivity of postsynaptic potentials and spike responses in cat visual cortex

Intracellular recordings were made from neurons in the cat visual cortex (area 17) to compare the orientation and direction selectivities of the output of a cell with those of the input the cell receives. The input to a cell was estimated from the PSPs (postsynaptic potentials) evoked by visual stimulation, and the output estimated from the number of spikes generated during the same responses. For the whole sample, selectivity of the output of cells was significantly higher than selectivity of their input. Upon PSP to spike transformation, the selectivity index was, on average, doubled. However, the degree of the selectivity improvement in individual cells was very different, varying from cases in which highly selective output was created from a poorly selective input and thus selectivity was greatly improved, to little or no improvement in other neurons. The improvement of selectivity was not correlated with resting membrane potential, threshold for action potential generation, background discharge rate or amplitude of optimal PSP response. Further, no systematic difference was found between simple and complex cells in the input-output relations, indicating that the 'tip of the iceberg' effect on shaping the response selectivity was cell specific, but not cell type specific. This supports the notion that multiple mechanisms are responsible for generation of the response selectivity, and that the contribution of any particular mechanism may vary from one cell to the other. The heterogeneity of the input-output relations in visual cortical cells could indicate different functions of cells in the cortical network; some cells are creating selectivity de novo, the function of other neurons probably being repetition and amplification of the selected signal and arrangement of the output of a whole column.     

Visual acuity of neurones in the cat lateral suprasylvian cortex

The spatial acuity was measured for cells of the posteromedial lateral suprasylvian area (PMLS) of the cat. Acuities were found to be 2 cycles/degree (15 min arc) at best, and 1 cycle/degree (30 min arc) on average. Both best acuity and average acuity remained constant with receptive field eccentricity within 20° of the area centralis, and then fell gradually with eccentricity. Acuity was good, given receptive field size, and was not correlated with receptive field size. Comparisons are drawn with other visual structures.     

Orientation selectivity of synaptic potentials in neurons of cat primary visual cortex

Neurons of the visual cortex of the cat were penetrated with intracellular electrodes and postsynaptic potentials evoked by visual stimuli recorded. By alternately polarizing the cell with steady current injected through the recording electrode, IPSPs and EPSPs could be recorded and analyzed independently. Hyperpolarizing current suppressed IPSPs and enhanced EPSPs by moving the membrane potential toward the IPSP equilibrium potential. Depolarizing the cell toward the EPSP equilibrium potential enhanced IPSP. The responses to electrical stimulation of the LGN, where EPSPs and IPSPs could be distinguished easily by virtue of their characteristic latencies and shapes, were used to set the current injection to the appropriate level to view the two types of synaptic potential. EPSPs were found to be well oriented in that maximal depolarizing responses could be evoked at only one stimulus orientation; rotating the stimulus orientation in either direction produced a fall in the EPSP response. IPSPs were also well tuned to orientation, and invariably the preferred orientations of EPSPs and IPSPs in any one cell were identical. In addition, no systematic difference in the width of tuning of the two types of potential was seen. This result has been obtained from penetrations of over 30 cortical cells, including those with simple and complex receptive fields. It is concluded that orientation of cortical receptive fields is neither created nor sharpened by inhibition between neurons with different orientation preference. The function of inhibition evoked simultaneously with excitation by optimally oriented stimuli has yet to be determined, though it is likely to be the mechanism underlying other cortical receptive field properties, such as direction selectivity and end-stopping.     

Parallel development of orientation maps and spatial frequency selectivity in cat visual cortex

In an early stage of the postnatal development of cats, orientation maps mature and spatial frequency selectivity is consolidated. To investigate the time course of orientation map maturation associated with the consolidation of spatial frequency selectivity, we performed optical imaging of intrinsic signals in areas 17 and 18 of cats under the stimulation of drifting square-wave gratings with different orientations and spatial frequencies. First, orientation maps for lower spatial frequencies emerged in the entire part of the lateral gyrus, which includes areas 17 and 18, and then these orientation maps in the posterior part of the lateral gyrus disappeared as orientation maps for higher spatial frequencies matured. Independent of age, an anteroposterior gradient of response strengths from lower to higher spatial frequencies was observed. This indicates that the regional distribution of spatial frequencies is innately determined. The size of iso-orientation domains tended to decrease as the stimulus spatial frequency increased at every age examined. In contrast, orientation representation bias changed with age. In cats younger than 3 months, the cardinal (vertical and horizontal) orientations were represented predominantly over the oblique orientations. However, in young adult cats from 3 to 9 months old, the representation bias switched to predominantly oblique orientations. These age-dependent changes in the orientation representation bias imply that orientation maps continue to elaborate within postnatal 1 year with the consolidation of spatial frequency selectivity. We conclude that both intrinsic and mutual factors lead to the development of orientation maps and spatial frequency selectivity.     

Sensory and multisensory representations within the cat rostral suprasylvian cortex

Because the posterior limb of the rostral suprasylvian sulcus (RSp) of the cat resides in close proximity to representations of the somatosensory, auditory, and visual modalities, the surrounding cortices would be expected to be a region where a high degree of multisensory convergence and integration is found. The present experiments tested this notion by using anatomical and electrophysiological methods. Tracer injections into somatosensory, auditory, and visual cortical areas almost all produced terminal labeling within the RSp, albeit at different locations and in different proportions. Inputs from somatosensory cortices primarily targeted the inner portion of the anterior RSp; inputs from auditory cortices generally filled the outer portion of the middle and posterior RSp; inputs from visual cortices terminated in the inner portion of the posterior RSp. These projections did not have sharp borders but often overlapped one another, thereby providing a substrate for multisensory convergence. Electrophysiological recordings confirmed this anatomical organization as well as identifying the presence of multisensory (bimodal) neurons in the areas of overlap between representations. Curiously, however, the proportion of bimodal neurons was only 24% of the neurons sampled in this region, and the majority of these did not show multisensory interactions when combined-modality stimuli were presented. In summary, these experiments indicate that the RSp is primarily auditory in nature, but this representation could be further subdivided into an outer sulcal anterior auditory field (sAAF) and an inner field of the rostral suprasylvian sulcus (FRS).     

The retinotopic organization of lateral suprasylvian visual areas in the cat

This is the second in a series of papers in which we describe our continuing efforts to define functional units of visual cortex based upon electro-physiological mapping of single and multiple unit activity in both awake and the nitrous oxide anesthetized cats. In the first paper (Tusa, Palmer and Rosenquist, '78), the extent and retinotopic organization of area 17 were described. In this paper, we describe the somewhat more complex organization of the visual cortex lying on the banks of the middle and posterior suprasylvian sulci. This region of cortex consists of six retinotopically organized units. These areas are arranged as three roughly mirror symmetrical pairs separated in each case by the fundus of the middle or posterior suprasylvian sulci. Some thalamo-cortical autoradiographic material is presented which supports this parcellation of the cortex.     

Effect of the richness of the environment on neurons in cat visual cortex. II. Spatial and temporal frequency characteristics

The quantitative distribution of synaptic contacts in the cat cerebral cortex is affected by the richness of the environment: the number of round-asymmetrical (RA) synapses per neuron is higher while the number of flat-symmetrical (FS) contacts is lower in the visual cortex of cats raised in an enriched environmental condition (EC) compared to those reared in an impoverished condition (IC). The size of FS synaptic contacts is also affected by the complexity of the animal's environment. It has been suggested that these morphological synaptic changes induced by the richness of the environment correlate with differences in physiological properties of the visual cortex. This question has been explored by assessing the cortical unit contrast sensitivity as a function of spatial and temporal frequency of area 17 cells in cats raised either in isolation (IC) or in a colony with ample environmental stimulation (EC). Contrast sensitivity is affected by the richness of the environment: at the preferred spatial frequency, an average enriched unit can detect lower contrasts (mean = 0.6%) than an average impoverished cell (0.9%; P less than 0.002). In addition, the mean highest spatial frequency that can be resolved (acuity) is greater in EC than in IC cells (1.7 and 1.2 cycles per degree, respectively, P less than 0.0001), whereas the spatial frequency at which units respond best is not statistically affected by the environment. The preferred temporal frequency also show a significant difference in EC (1.0-6.5 Hz) and IC units (0.9-4.0 Hz; P less than 0.01).(ABSTRACT TRUNCATED AT 250 WORDS)     

Spatial and temporal characteristics of visual motion perception involving V5 visual cortex

The anatomical substrates of the perception of motion have not yet been established in a detailed way on an individual level. The aim of this study was to develop a systematic procedure for mapping the visual cortex using Transcranial Magnetic Stimulation (TMS). The results showed that such an individual and detailed map of the spatial and temporal characteristics of motion perception can be constructed using TMS.     

Spatial and temporal characteristics of visual motion perception involving V5 visual cortex

Abstract

The anatomical substrates of the perception of motion have not yet been established in a detailed way on an individual level. The aim of this study was to develop a systematic procedure for mapping the visual cortex using Transcranial Magnetic Stimulation (TMS). The results showed that such an individual and detailed map of the spatial and temporal characteristics of motion perception can be constructed using TMS. [Neurol Res 2002; 24: 266-270]     

Spatial and temporal distribution of visual information coding in lateral prefrontal cortex

Prefrontal neurons code many kinds of behaviourally relevant visual information. In behaving monkeys, we used a cued target detection task to address coding of objects, behavioural categories and spatial locations, examining the temporal evolution of neural activity across dorsal and ventral regions of the lateral prefrontal cortex (encompassing parts of areas 9, 46, 45A and 8A), and across the two cerebral hemispheres. Within each hemisphere there was little evidence for regional specialisation, with neurons in dorsal and ventral regions showing closely similar patterns of selectivity for objects, categories and locations. For a stimulus in either visual field, however, there was a strong and temporally specific difference in response in the two cerebral hemispheres. In the first part of the visual response (50–250 ms from stimulus onset), processing in each hemisphere was largely restricted to contralateral stimuli, with strong responses to such stimuli, and selectivity for both object and category. Later (300–500 ms), responses to ipsilateral stimuli also appeared, many cells now responding more strongly to ipsilateral than to contralateral stimuli, and many showing selectivity for category. Activity on error trials showed that late activity in both hemispheres reflected the animal's final decision. As information is processed towards a behavioural decision, its encoding spreads to encompass large, bilateral regions of prefrontal cortex.     

Acoustic tuning of single cells in middle suprasylvian cortex of cat

Acoustic tuning of single cells in medial suprasylvian ‘association’ cortex (MSA) was studied in cats anesthetized with α-chloralose and in locally anesthetized cats immobilized with Flaxedil. MSA cells in the chloralose preparations exhibited little or no spontaneous activity and onset responses of variable latency to tone pulse stimuli. Tuning curves for these cells were extremely broad (extending over 4–5 octaves or more) and insensitive (thresholds of 40–60 dB SPL). Cells recorded from AI in the same preparations exhibited the sharp tuning and high sensitivity reported by others. In the immobilized animals, spontaneous firing rates of MSA cells were much higher and responses to tones were poorly defined and extremely labile. Thresholds could not easily be determined for many of these cells, but in some cases it was possible to establish approximate response areas. These cells exhibited a breadth of response and lack of sensitivity comparable to that seen under chloralose. The data indicate a high degree of convergence in the acoustic input to cells in MSA. Although inter-modality comparison is difficult, acoustic input to this area appears to be less specifically organized than visual input.     

Action selectivity in parietal and temporal cortex

The sensory-action theory proposes that the neural substrates underlying action representations are related to a visuomotor action system encompassing the left ventral premotor cortex, the anterior intraparietal (AIP) and left posterior middle temporal gyrus (LPMT). Using fMRI, we demonstrate that semantic decisions on action, relative to non-action words, increased activation in the left AIP and LPMT irrespective of whether the words were presented in a written or spoken form. Left AIP and LPMT might thus play the role of amodal semantic regions that can be activated via auditory as well as visual input. Left AIP and LPMT did not distinguish between different types of actions such as hand actions and whole body movements, although a right STS region responded selectively to whole body movements.