Phase of responses to sinusoidal gratings of simple cells in cat striate cortex.
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Biomedical subjects
Publications and source records attributed to B B Lee.
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The responses of neurones in the lateral geniculate nucleus (LGN) were investigated in anaesthetised rhesus monkeys. A new classification for cells in the parvocellular layers (PCL) is proposed, based on their spectral response curve and their response to white stimuli: (A) narrow-band, short wavelength (NS) excited cells, activity suppressed by white stimuli; (B) wide-band, short-wavelength (WS) excited cells, excited by white stimuli; (C) wide-band, long-wavelength (WL) excited cells, (D) narrow-band, long-wavelength (NL) excited cells, activity suppressed by white stimuli; (E) light suppressed (LI) cells, activity suppressed by all wavelengths, usually with some concealed excitatory input at extreme short or long wavelengths. Responses to moving bars and to spots of various diameters (area response curves) were determined for various wavelengths. It was found that the receptive fields from which wavelength-dependent excitatory or suppressive effects could be elicited are concentrically superimposed. The spectral responsiveness of the excitatory inputs to individual cell types corresponds to the absorption curves of single cones (S-, M- or L-cone for NS, WS and WL cells respectively), the spectral distribution of the suppressive mechanisms of all cells was panchromatic and approximately fitted to a sum of all cones. The excitatory input to NL-cells cannot be related to any of the known cone absorption curves, and a simple (L-M) subtraction model is questioned. Neurones in the magnocellular layers (MCL) can be divided into on- and off-centre cells as in the cat's LGN and give qualitatively similar responses over the whole spectrum. In contrast to the tonic responses of PCL cells, MCL cells respond phasically to chromatic and white flashed spots, even with the smallest stimuli. Implications of these findings for colour processing in the LGN are discussed.
The responses to moving stimuli of single cells in the parvo- and magnocellular layers (PCL and MCL) of the macaque lateral geniculate nucleus (LGN) have been studied. PCL cells respond with a monophasic increase or decrease in firing when a bar passes across the receptive field, according to the wavelength composition of the stimulus. MCL cells respond with a biphasic sequence of excitation and suppression or vice versa dependent on whether a cell is on-centre or off-centre and on stimulus contrast direction. With large stimuli, PCL cells respond as long as the stimulus covers the receptive field while MCL cells respond only at the contrast borders. MCL cell responses are maximal with bars just long enough to cover the field centre, while PCL cell responses show a variable relation with bar length, depending on stimulus wavelength and receptive field structure. PCL cells show broad velocity tuning while at least some MCL cells were more sharply tuned. Many cells in the macaque LGN show weak orientation or direction preference.
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The activity of retinal ganglion cells and cortical cells with overlapping receptive fields was simultaneously recorded. The responses to moving stimuli of individual simple cortical cells could be accounted for on the basis of the cell receiving either on-centre or off-centre afferents; instances in which it was necessary to postulate a mixed on- and off-centre input were not found. In six instances cross correlograms of ganglion cell and cortical cell activity showed that the ganglion cell was afferent, via a relay cell in the LGN, to the cortical neurone. The receptive fields of such pairs were almost completely overlapping and concentric. In three cases a sustained ganglion cell projected to a simple cortical cell. In one case a transient ganglion cell projected to a simple cell, and in one case a sustained and a transient ganglion cell projected to the same simple cell.
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.
The receptive fields of LGN cells were investigated with stationary light and dark spot and annulus stimuli. Stimulus size and background intensity were varied while stimulus/background contrast was kept constant. The speed of dark adaptation vaired considerably from cell to cell. Dark adaptation made responses more sustained in all neurones and eliminated the oscillatory on-responses evoked under some conditions in the light-adapted cells. Dark adaptation led also to a disappearance of early phasic inhibition in on-responses, and increased response rise time and latency. The power of surround responses to inhibit centre responses decreased slightly at low levels of light adaptation in LGN cells but much less than in retinal ganglion cells. Some other traces of changing retinal surround effects also appeared inthe LGN on dark adaptation. For example, the functional size of receptive fields increased at low levels of illuminance as has been observed in retinal ganglion cells and the receptive fields as estimated from response peaks were larger than those estimated from sustained components.
The responses of neurones in laminae A and A1 of the cat lateral geniculate nucleus to moving stimuli were investigated at different background luminances. Moving bright slits, dark bars and edges were employed; the contrast of stimuli against the background was held constant. Background intensities varied from 10(-3) to 10(2) td. Responses as stimuli passed across the centres of LGN receptive fields became stronger with increasing levels of light adaptation up to 10(-1)-10(1) td and then remained constant. Responses as stimuli passed through surround regions altered qualitatively with adaptation level, generally increasing in strength and complexity with background luminance. As a bright slit for on-centre cells or dark bar for off-centre cells left the surround, in almost all units a strong secondary peak could be elicited by an appropriate selection of the adaptation conditions. Many features of the responses to moving stimuli could not be predicted from the responses to stationary stimuli under different adaptation conditions described in the previous paper.
Macaques were trained to respond to electrical excitation applied through electrodes permanently implanted within or upon striate cortex. Threshold current for the animal to detect this stimulation was highly consistent from day to day and, in the absence of tissue encapsulation of the electrodes or deliberately inflicted damage, remained stable indefinitely, 38 months in the longest case so far. Stimulating continuously for 1-8h, however, produces an elevation of threshold, which may be permanent or temporary, depending upon a variety of conditions. A major cause of such injury is the hydrolysis commonly occurring consequent to passage of low-level currents between solutions and metal electrodes. Even when the hydrolytic reaction is eliminated by restricting the level of electrode polarization or by using capacitative stimulation with tantalum pentoxide electrodes, a rise in threshold often still occurs with protracted stimulation. With proper control in some instances, however, effective stimulation at 2-10 times the threshold level could be maintained indefinitely without apparent injury, e.g. in a blind monkey having a threshold of 290 muA that could respond immediately to an 80-muA diminution in 580-muA, 0.2-msec stimulus pulses which had been applied steadily for 1 h at 50 Hz.
Using permanently implanted electrodes in squirrel monkeys and macaques, transmission through the lateral geniculate nucleus (LGN) was assayed from the amplitude of potentials evoked in optic radiation by and electrical pulse applied to optic tract. Averaging of either individually or machine selected potentials, elicited at 0.3, 1.0, 20 or 50 HZ, in all cases showed a decrease in transmission ranging from 5-60% in the period after saccadic eye movements made ad libitum. The suppression was greater in a patterned visual environment than in diffuse illumination, which in turn was greater than that occurring following saccades in the dark. Demonstration of the effect in darkness always required data averaging and never exceeded 20%. The effect was consistently greater in the magnocellular than parvocellular component. Suppresion was often abruptly terminated and replaced by a facilitation of 5-15% about 100 msec after saccade detection. Comparable effects were observed for excitability of striate cortex tested by a stimulus pulse applied to optic radiation. In addition, sharply demarcated potentials inherently arising in LGN and striate cortex were found in association with saccades made even in total darkness. Neglecting a possible but dubious contribution from eye muscle proprioceptors, the experiments establish the existence of a centrally originating modulation of visual processing at both LGN and striate cortex in ralation to saccadic eye movement in primates. This modulation may partially underlie the phenomenon of "saccadic suppression" and hasten the acquistion of a meaningful visualsample immediately following an ocular saccade. It remains uncertain as to how it may relate to similar or greater effects accompanying changes in alertness, or to fluctuations of unknown origin occurring sometimes semirhythmically at 0.05-0.03 HZ (Fig 7).
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