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P Heggelund

Publications and source records attributed to P Heggelund.

At least 19 recordsLinked to original sources
PubMed1992

A bidimensional theory of achromatic color vision.

The theory is based on a perceptive color system where the achromatic colors are specified by their degree of similarity to the three qualities white, black and luminous. Black and luminous are treated as opponent variables. It is assumed that white and luminous/black are determined by different kinds of visual processes termed the w- and the b-process. The relationship between these processes and luminance parameters in a simple disc/ring configuration is derived from available data. The b-process is related to stimulus contrast in a simple manner. It is assumed to involve cells with antagonistic center/surround organization of the receptive field. The w-process is primarily determined by the local luminance, and it is assumed to involve cells that lack a center/surround organization of the receptive field. The w-process has properties similar to the processes involved in chromatic color vision. The theory can account for different kinds of psychophysical data on achromatic colors like data on simultaneous contrast, color scaling, and color constancy.

Color Perception↗
PubMed1992

The effect of contrast on the visual response of lagged and nonlagged cells in the cat lateral geniculate nucleus.

The response vs. contrast characteristics of different cell classes in the dorsal lateral geniculate nucleus (LGN) were compared. The luminance of a stationary flashing light spot was varied stepwise while the background luminance was constant. Lagged X cells had lower slope of the response vs. contrast curve (contrast gain), and they reached the midpoint of the response range over which the cells' response varied (dynamic response range) at higher contrast than nonlagged X cells. These results indicated that nonlagged cells are well suited for detection of small contrasts, whereas lagged cells may discriminate between contrasts over a larger range. The contrast gain and the contrast corresponding to the midpoint of the dynamic response range were similar for X and Y cells. The latency to onset and to half-rise of the visual response decreased with increasing contrast, most pronounced for lagged cells. Even at the highest contrasts, the latency of lagged cells remained longer than for nonlagged cells. For many lagged cells, the latency to half-fall decreased with increasing contrast. It is shown that the differences in the response vs. contrast characteristics between lagged and nonlagged X cells in the cat are similar to the differences between the parvocellular and magnocellular neurones in the monkey.

Animals↗
PubMed1990

Neurotransmitter receptors mediating excitatory input to cells in the cat lateral geniculate nucleus. I. Lagged cells.

1. Synaptic mechanisms that might explain the functional properties of the recently discovered class of lagged cells in the dorsal lateral geniculate nucleus (LGN) were analyzed with electrophysiological and pharmacologic techniques. To study the type of excitatory amino acid (EAA) receptor that mediates visual responses of lagged cells, we recorded the response of single cells to a stationary flashing light spot before, during, and after microiontophoretic application of antagonists and agonists to EAA receptors. 2. The visual response of the lagged cells could be almost completely blocked by an antagonist to the N-methyl-D-aspartate (NMDA) receptor. The degree of suppression was dose dependent, and the average maximum degree of suppression for all the cells was 94%. NMDA enhanced the response, and this enhancement was antagonized by NMDA antagonists. A quisqualate/kainate receptor antagonist had no significant effect on the lagged cells. 3. These findings indicate that the visual response in lagged cells is dependent upon activation of NMDA receptors, which may directly result from activation of retinal inputs. 4. No pharmacologic difference was seen between lagged X- and Y-cells, or between lagged ON- and OFF-center cells. 5. gamma-Aminobutyric acid-A (GABA-A) receptor antagonists were used to study whether the characteristic lag of the visual response and the suppression of the initial transient response component of the lagged cells are dependent on geniculate inhibition. Beside enhancement of the visual response, the GABA antagonists strongly reduced the lag of the visual response, and an initial transient response component occurred instead of the initial suppression. The lag remained slightly longer than for nonlagged cells, and the peak firing rate of the transient was below values typical for nonlagged cells, indicating that the lagged cell properties are dependent on other factors beside GABA-A receptor-mediated inhibition. 6. The enhanced visual response during iontophoresis of GABA antagonists could be completely blocked by simultaneous iontophoresis of an NMDA-receptor antagonist. This gives further support to the hypothesis that the retinal input to these cells is mediated by NMDA receptors. 7. The NMDA-receptor/ionophore complex mediates excitatory postsynaptic potentials (EPSPs) characterized by slow rise and decay times and long duration. The ionophore is characterized by a voltage-dependent blockade that makes these receptors particularly sensitive to inhibitory input. The temporal interplay between the slow NMDA receptor-mediated EPSPs and the fast GABA receptor-mediated inhibitory postsynaptic potentials (IPSPs) may explain the characteristic response properties of the lagged cells.

Afferent Pathways↗
PubMed1990

Neurotransmitter receptors mediating excitatory input to cells in the cat lateral geniculate nucleus. II. Nonlagged cells.

1. We studied the type of receptor for excitatory amino acids (EAA) that mediates visual responses of nonlagged cells in the dorsal lateral geniculate nucleus (LGN) by recording the visual response of single cells to a stationary flashing spot before, during, and after iontophoretical application of antagonists and agonists to EAA receptors. 2. The visual response of the nonlagged cells was strongly suppressed, in a dose-dependent manner, by the specific quisqualate/kainate receptor antagonist 6,7-dinitroquinoxaline-2,3-dione (DNQX). The average degree of suppression was 74%. Quisqualate enhanced the visual response. 3. Specific antagonists to the N-methyl-D-aspartate (NMDA) receptor had a weak suppressing effect on most nonlagged cells. The average degree of suppression was 22%. Measurement of such weak effects was complicated by the considerable spontaneous fluctuations of responsivity in the LGN cells. In the majority of cells where the visual response was suppressed by NMDA antagonists, the tonic response component was more strongly suppressed than the initial transient response component. The visual response was enhanced by NMDA, and this enhancement was antagonized by NMDA antagonists. 4. These findings suggest that the excitatory input from the retina to nonlagged LGN cells is mainly mediated by non-NMDA receptors. The non-NMDA receptors mediate fast EPSPs, and this can explain the fast onset and offset of the visual response of the nonlagged cells. 5. The generally small contribution from NMDA receptors to the visual response of the nonlagged cells might reflect a minor involvement of these receptors in the retinal input, or it could be related to the excitatory input to LGN from the visual cortex. 6. To study whether the expression of NMDA receptors was related to modulatory brain stem input to LGN, we examined the effects of the NMDA antagonists when the visual response was enhanced with gamma-aminobutyric acid (GABA) antagonists or acetylcholine (ACh). Neither of these pharmacologic manipulations consistently increased the relative contribution of NMDA receptors to the visual response. 7. No pharmacologic difference was found between nonlagged X- and Y-cells, or between ON- and OFF-center cells.(ABSTRACT TRUNCATED AT 400 WORDS)

Afferent Pathways↗
PubMed1989

Response to rates of luminance change of sustained and transient cells in the cat lateral geniculate nucleus and optic tract.

We recorded the response of sustained (X) and transient (Y) cells in the cat lateral geniculate nucleus (LGN) and optic tract to a stationary spot while the spot luminance was increased and decreased with a constant rate (linear luminance functions), or modulated sinusoidally. The spot filled the receptive field center, and was surrounded by an annulus of fixed luminance. The LGN X cells seemed to perform a differentiation-like operation in the time domain at slow temporal modulations, giving information about rate of luminance change. To the linear luminance functions the cells responded with a constant firing rate. The on-center cells were activated during increasing luminance, the off-center cells during decreasing luminance. This firing rate increased monotonically with rate of luminance change. To low-frequency sinusoidal modulations the cells had a marked negative phase shift. The response of the LGN Y cells had a transient component shortly after the luminance started to increase (on-center cells) or decrease (off-center cells), followed by a secondary, gradually changing component. The peak of the transient component occurred on average when the response of the X cells increased most rapidly. To low-frequency sinusoidal modulation the average negative phase shift of this peak was twice the average of the X cells. The Y system could accordingly provide information about rate of change in the response of the X system. In the optic tract the X fiber response resembled the LGN X cell response in most respects. The Y fibers had only a weak transient response component, so this component was accentuated in the thalamic relay. Also the sensitivity for rate of luminance change was increased in LGN.

Action Potentials↗
PubMed1989

Postnatal development of glutamatergic, GABAergic, and cholinergic neurotransmitter phenotypes in the visual cortex, lateral geniculate nucleus, pulvinar, and superior colliculus in cats.

We have analyzed the postnatal development of glutamatergic/aspartergic, GABAergic, and cholinergic neurotransmitter systems in the visual cortical Areas 17 and 18, lateral geniculate nucleus (LGN), pulvinar, and the visual and non-visual parts of superior colliculus (SC) in kittens. High-affinity uptake of D-aspartate (HA D-Asp), glutamate decarboxylase (GAD), and choline acetyltransferase (ChAT) activities were measured as a means of probing the development of the respective transmitter systems. HA D-Asp exceeded the adult level several-fold in all areas during the postnatal period which corresponded with the period of maximal dendritic/axonal branching patterns and synapse densities in the respective regions. GAD exhibited a gradual increase towards adult levels during the first month. The adult level was reached during postnatal week (PNW) 5-6 in Areas 17 and 18, during PNW3 within LGN, pulvinar, and the visual part of SC. In the nonvisual part of SC, the adult GAD level was reached as early as PNW2. ChAT exhibited biphasic developmental profiles in Areas 17 and 18. An initial peak of near adultlike activity in PNW2 was followed by a decline and subsequently by a slow increase towards adult levels during PNW5-17. ChAT developed very slowly in LGN and pulvinar, and in the latter structure only approximately 70% of the adult activity had been attained by PNW17. In both subdivisions of SC, ChAT had reached adult levels during PNW3-5. Dark-rearing from birth until PNW6 moderately attenuated GAD development in all areas and increased ChAT activity in Areas 17 and 18 but did not affect development of HA D-Asp in any part of the kitten visual system. Our neurochemical findings in the developing cat visual system are consistent with available evidence regarding localization of neurotransmitter systems, as well as postnatal changes in terms of cytoarchitectonics, synaptogenesis, functional development, and susceptibility to neonatal dark-rearing in visual pathways.

Animals↗
PubMed1987

Reduced binocularity in the noradrenaline-infused striate cortex of acutely anesthetized and paralyzed, otherwise normal cats.

In anesthetized and paralyzed cats, the normal alignment of the visual axes is disturbed by paralysis of the eye muscles. Thus, the separation between paired receptive fields of binocular cells in visual cortex is increased (paralysis squint). This increased separation is normally tolerated by the majority of visuocortical cells, about 80% of them being binocularly driven (Hubel and Wiesel 1962). It was shown previously that neuronal plasticity in visual cortex can be enhanced in both normal adult cats (Kasamatsu et al. 1979) and kittens (Kuppermann and Kasamatsu 1984) by intracortical microinfusion of noradrenaline (NA). In the present study we tested whether the usual range of disparity produced by the paralysis squint is sufficient to induce ocular dominance changes in visual cortex of adult cats when the neuronal plasticity is enhanced by NA. NA was continuously infused into visual cortex throughout the experiments. The period of the paralysis squint varied from experiment to experiment between 9 and 47 h. We found: (1) These short periods were sufficient to produce a marked reduction in the proportion of binocular cells. (2) The proportion decreased linearly with increasing the duration of the squint period at a rate of 0.17 per 10 h up to about 22 h. (3) At longer durations the average binocularity remained at about 0.30 and could not be further reduced in the present paradigm. (4) The binocularity seemed to decrease with increasing separation of paired receptive fields. (5) Binocularity increased again toward the normal value after optical correction of the squint. (6) The amount of increased binocularity was linearly correlated with the duration of the period after the squint correction. (7) The binocularity increased at a rate of 0.18 per 10 h, reaching the normal value in less than 30 h. We thus concluded that if visuocortical plasticity is maintained at a high level through the continuous infusion of NA it is possible to change the ocular dominance distribution in the mature visual cortex by manipulations of the alignment of the visual axes even in the acutely anesthetized and paralyzed condition.

Anesthesia↗
PubMed1986

Quantitative studies of the discharge fields of single cells in cat striate cortex.

The configuration and width of on- and off-response zones in the discharge field of single cells in cat striate cortex was analysed by quantitative methods. The responses across on- and off-zones were plotted for 321 cells with a stationary optimum oriented light slit. The cells fell into two completely distinct subgroups with respect to the degree of overlap between adjacent on- and off-zones. The simple cells had a mean overlap of 16.8%, the complex cells 94.5%. For simple cells the ratio between the maximum off- and maximum on-response in the discharge field was bimodal, showing that two distinct subgroups termed on- and off-dominant cells could be distinguished. For the complex cells the corresponding frequency distribution was unimodal. The maximum response on the two regions adjacent to the most responsive discharge zone (the dominant zone) differed markedly for most simple cells, and only a very few cells had discharge fields approximating an ideal even symmetric field. The frequency distribution of the ratio between the maximum response in the two regions was unimodal showing that odd and even symmetric fields did not form distinct subgroups of simple cells. The number of different discharge zones in simple cells varied from one to five. The zones were arranged as alternating on- and off-zones across the discharge field. The maximum response in the subzones decreased with increasing sequential distance from the dominant zone, so the response pattern across each side of the discharge field resembled a damped wave-form pattern. All the complex cells had one on- and off-zone which overlapped. The mean width of the subregions in the simple cell discharge field and the mean distance between the response maxima in the subzones increased in the same proportion with increasing eccentricity. The paracentral fields were therefore like magnified central fields. The average width of the whole discharge field was not significantly different for the simple and the complex cells at the various eccentricities.

Action Potentials↗
PubMed1986

Quantitative studies of enhancement and suppression zones in the receptive field of simple cells in cat striate cortex.

The configuration and extension of enhancement and suppression zones were compared with the configuration and extension of on- and off-response zones across the receptive field of simple cells in cat striate cortex. The enhancement and suppression zones were determined by a dual-stimulus technique where a stationary flashing light slit produced activity against which activation profiles across the receptive field were plotted with a parallel stationary test slit. The activation profiles showed less variation in receptive field configuration than plots of on- and off-discharge zones. Whereas the number of on- and off-zones varied between one and five, the activation profiles showed at least three distinct subregions in the receptive field, i.e. a central zone with an adjacent oppositely responding zone on each side. The responsivity was clearly stronger in one of these proximal flank zones. An additional zone occurred distal to the strong proximal flank zone in 53% of the cells, and in 10% such a distal zone occurred distal to both proximal flank zones. There was good correspondence between the location of on- and off-discharge zones and the location of the enhancement and suppression regions, although some subregions seen in the activation profiles did not appear in plots made with a single slit. The maximum discharge and the maximum enhancement and suppression effects in a subregion were found at the same receptive field location. The width of a subregion was measured as the width of the eventual on- or off-discharge zone determined with a single slit, as the width of the enhancement zone, and as the width of the suppression zone determined with the dual-slit technique. The enhancement zone was narrower, and the suppression zone wider than the discharge zone. The strong proximal flank zone had the same width as the central zone, but was wider than the weak proximal flank zone. For most cells the distances between successive extreme points across the activation profiles were constant, and this may explain the selectivity of the cells for spatial frequency of periodic stimuli. The strongest flank suppression occurred for most cells in that of the two proximal flank zones which had the strongest discharge to the single slit. Nevertheless, there was no correlation between the degree of discharge and the degree of suppression produced by opposite light cycles.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1985

Plasticity in cat visual cortex restored by electrical stimulation of the locus coeruleus.

It has been proposed that the presence of noradrenaline (NA)-containing terminals and NA-related receptors within the visual cortex is necessary to maintain the high level of neuronal plasticity in the immature visual cortex of kittens. In the present study we wanted to show whether electrical stimulation of the locus coeruleus (LC), which contains the somata of these cortical NA fibers, can restore neuronal plasticity to the normally aplastic visual cortex of juvenile and adult cats. We consistently found a significant loss of binocular cells in the visual cortex of mature animals which had monocular vision for only 12 h dispersed over 6 days (2 h a day, otherwise kept in the dark) in combination with concurrent LC stimulation. This result was interpreted as indicating that endogenous NA released from NA terminals restored susceptibility to monocular vision in the mature visual cortex. We next examined how long the restored plasticity lasts in the same animals after the LC stimulation was ended. The animals revived from the first recording session were either returned to the same daily schedule of brief monocular exposure (light/dark = 2/22 h) as before, or subjected to the usual monocular lid suture and kept in a cat colony environment (light/dark = 16/8 h). The LC electrodes had been removed and no more electrical stimulation was delivered at this stage. In the animals subjected to reiteration of brief monocular exposure, the state of reduced binocularity gradually returned to normal over a period of 2-3 weeks after stopping LC stimulation. We calculated that the revived plasticity disappeared at an average rate of a 22% loss every 7 days. This result sharply contrasted with the result obtained in the animals subjected to usual monocular lid suture. In this test the state of reduced binocularity continued for at least the next 3 weeks, suggesting that the restored plasticity was sustained throughout a period of 3 weeks (longest term tested). The different results obtained in the two paradigms may be explained by the different strength of binocular imbalance in the two tests imposed on the visual cortex in which neuronal plasticity was restored partially.

Animals↗
PubMed1985

Development of spatial receptive-field organization and orientation selectivity in kitten striate cortex.

The functional organization of the receptive field of neurons in striate cortex of kittens from 8 days to 3 mo of age was studied by extracellular recordings. A quantitative dual-stimulus technique was used, which allowed for analysis of both enhancement and suppression zones in the receptive field. Furthermore the development of orientation selectivity was studied quantitatively in the same cells. Already in the youngest kittens the receptive fields were spatially organized like adult fields, with a central zone and adjacent flanks that responded in opposite manner to the light stimulus. The relative suppression in the subzones was as strong as in adult cells. Both simple and complex cells were found from 8 days. The receptive fields were like magnified adult fields. The width of the dominant discharge-field zone and the distance between the positions giving maximum discharge and maximum suppression decreased with age in the same proportions. The decrease could be explained by a corresponding decrease of the receptive-field-center size of retinal ganglion cells. Forty percent of the cells were orientation selective before 2 wk, and the fraction increased to 94% at 4 wk. Cells whose responses could be attenuated to at least half of the maximal response by changes of slit orientation were termed orientation selective. The half-width of the orientation-tuning curves narrowed during the first 5 wk, and this change was most marked in simple cells. The ability of the cells to discriminate between orientations in statistical terms was weak in the youngest kittens due to a large response variability, and showed a more pronounced development than the half-width did. The orientation-tuning curves were fitted by an exponential function, which showed the shape to be adultlike in all age groups. Two kittens were dark reared until recording at 1 mo of age. The spatial receptive-field organization and the orientation selectivity in these kittens were similar to normal-reared kittens at 1 mo. The responsivity of the cells of the dark-reared kittens was lower, and the latency before firing was longer than in the normal-reared kittens of the same age, and these response properties were more similar to those in 1- to 2-wk-old normal kittens. Our results indicate that the spatial organization of the receptive field is innate in most cells and that visual experience is unnecessary for the organization to be maintained and for the receptive-field width to mature during the first month postnatally.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1978

Response variability and orientation discrimination of single cells in striate cortex of cat.

The response of single cells in the striate cortex of cat to a moving light bar of variable orientation was measured by a method providing data on the mean response as well as the standard deviation (SD) at the different stimulus orientations. At the optimal stimulus orientation the SD was about 1/3 of the mean response. Marked differences in this respect were found between simple and complex cells, i.e., the SD for the simple cells was about 1/2 of the mean response and about 1/4 for the complex cells. The variation coefficient (Vc = SD/mean) was minimal at the optimal orientation and increased relatively in the same manner for simple and complex cells as the stimulus orientation was varied away from optimal orientation. The Vc varied with the mean response at optimal orientation in a nonlinear manner. A function is proposed which fits this relationship and which is equally applicable for both simple and complex cells. The mean orientation discrimination (MOD) was defined as that change in orientation angle away from the optimal which produced a response statistically different--on the 1% level--from the response to the optimal orientation. There were differences in MOD between the two sides of the orientation tuning curve: the mean of the smaller of the two values was 13.5 deg and of the larger 19.7 deg. No significant difference in MOD was found between simple and complex cells despite the fact that the halfwidth of the tuning curves for the two cell types was 19.5 deg and 31.6 deg, respectively. The preciseness in localization of the most sensitive part within the receptive field of single cells was calculated from the variability in time of occurrence of the smallest interspike interval. The degree of preciseness was found to be of the order of 1/4 of the receptive field diameter in both simple and complex cells. When nonoptimal stimulus orientations were presented, the preciseness significantly decreased in complex cells whereas it remained unchanged in simple cells. It is suggested that the same type of intracortical wiring produces orientation selectivity in simple and complex cells, and that the differences in tuning width are mainly due to a larger extension of inhibitory fields in the simple cells. Considering the cortical visual cells as elementary units in a network built for orientation detection and discrimination, the tuning width seems of minor importance for that function.

Animals↗
PubMed1977

The depth distribution of optimal stimulus orientations for neurones in cat area 17.

Neurones recording during penetrations through cat area 17 as near parallel to the radial fibre bundles as possible have been quantitatively tested as to their optimal orientation. Optimal orientation within any one penetration was similar though considerable variability was observed. Histological reconstruction and other considerations showed that this variability could not be attributed to poor penetration angle or limitations of the microelectrode technique. These results confirm that neurones with similar optimal orientations are found in all cortical layers at one cortical locus, but it is difficult to reconcile the variability observed with a mosaic-like distribution of orientation across the cortical surface. The findings were consistent, however, with the assumption of a continuous distribution of orientation sensitivity across the cortical surface with considerable superimposed scatter.

Animals↗
PubMed1975

Responses of striate cortical cells to moving edges of different curvatures.

The responses of twenty cells to stimuli of varying curvature were measured in cat's striate cortex. All the investigated cells were sensitive to the orientation of lines and not hypercomplex. Fourteen cells showed a systematic change of response with curvature. The optimal curvatures of the cells were distributed over the whole range investigated. Six cells were insensitive to curvature. The responses from all the typical simple cells (8) varied with curvature, whereas all the complex cells (5) were insensitive to curvature changes. The curvature tuning curves were broad and the variability to individual stimuli was high, independent on whether the cell responded best to straight or to curved edges. The findings do not support the view that individual cells of area 17 could detect curvature.

Animals↗
PubMed1975

Neural plasticity in visual cortex of adult cats after exposure to visual patterns.

Over a period of 2 weeks, adult cats were twice a day exposed for 1 hour to a visual environment consisting only of vertical stripes and for the rest of the time were kept in darkness. Subsequent investigation of the striate cortex showed a decrease in the number of neurons sensitive to orientations around the vertical relative to those sensitive to horizontal orientations. This indicates that plasticity of functional properties of the cortical neuronal network still exists in adult animals.

Animals↗