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A B Bonds

Publications and source records attributed to A B Bonds.

47 records · Page 3Linked to original sources

Image quality of the cat eye measured during retinal ganglion cell experiments.

1. The modulation transfer function (MTF) of the dioptrics of fifteen cat eyes was determined. The aerial image, formed by the eye of a standard object (a 0.5-1.0 degrees annulus), was photographed. The transmission of the film negative was measured with a scanning microdensitometer to yield the light distribution within the aerial image. Correcting for the double passage, this experimentally determined light distribution and the known object light distribution were used to obtain the MTF, applying Fourier methods. Each MTF was used to calculate the light distribution within the retinal image of stimuli of various geometry used in experiments on retinal ganglion cells in the same eye.2. When the eye was equipped with an artificial pupil of the same size as that used in the neurophysiological experiments (4.0-4.8 mm diam.) the MTF had fallen to 0.5 at 2.43 c/deg. When the pupil was removed the MTF had fallen to 0.5 at a much lower spatial frequency (1.0 c/deg). This shows that even when one uses an artificial pupil too large to provide optimal image quality there is a vast improvement over using no pupil.3. These image quality measurements were prompted by the need to know the actual stimulus image in experiments on the functional organization of the receptive field, a need exemplified in this paper by a few specific physiological results. The full neurophysiological results appear in the next two papers.

Action Potentials↗

Two-dimensional receptive-field organization in striate cortical neurons of the cat.

The two-dimensional organization of receptive fields (RFs) of 44 cells in the cat visual cortex and four cells from the cat LGN was measured by stimulation with either dots or bars of light. The light bars were presented in different positions and orientations centered on the RFs. The RFs found were arbitrarily divided into four general types: Punctate, resembling DOG filters (11%); those resembling Gabor filters (9%); elongate (36%); and multipeaked-type (44%). Elongate RFs, usually found in simple cells, could show more than one excitatory band or bifurcation of excitatory regions. Although regions inhibitory to a given stimulus transition (e.g. ON) often coincided with regions excitatory to the opposite transition (e.g. OFF), this was by no means the rule. Measurements were highly repeatable and stable over periods of at least 1 h. A comparison between measurements made with dots and with bars showed reasonable matches in about 40% of the cases. In general, bar-based measurements revealed larger RFs with more structure, especially with respect to inhibitory regions. Inactivation of lower cortical layers (V-VI) by local GABA injection was found to reduce sharpness of detail and to increase both receptive-field size and noise in upper layer cells, suggesting vertically organized RF mechanisms. Across the population, some cells bore close resemblance to theoretically proposed filters, while others had a complexity that was clearly not generalizable, to the extent that they seemed more suited to detection of specific structures. We would speculate that the broadly varying forms of cat cortical receptive fields result from developmental processes akin to those that form ocular-dominance columns, but on a smaller scale.

Algorithms↗

The influence of input from the lower cortical layers on the orientation tuning of upper layer V1 cells in a primate.

The receptive fields of cells in the primary visual cortex (area 17 or V1) show clear orientation selectivity, unlike those of the lateral geniculate nucleus (LGN) cells which provide their visual input. The intrinsic circuitry of V1 cells is believed to be partly responsible for this selectivity. We investigated the influence of ascending projections from neurons in the lower layers (5 and 6) of V1 on the orientation selectivity of single neurons in the upper layers (2,3, and 4) by reversibly inactivating ("blocking") lower layer neural activity with iontophoretic application of gamma-aminobutyric acid (GABA) while recording from upper layer cells in the prosimian primate, Galago crassicaudatus. During lower layer blocking, the majority (20/28 = 71.4%) of upper layer neurons exhibited a change in the orientation of their preferred stimulus, a reduction in their orientation tuning, and/or an increase in their response amplitude. Twelve (42.9%) neurons exhibited shifts in their preferred orientation averaging 11 (+/- 4) deg. These neurons were located on average, 272 (+/- 120) microns tangential from the vertical axis of the pipette center. Eleven neurons (39.2%) exhibited an average reduced orientation tuning of 52.5%. Their average location was 230 +/- (115) microns away from the vertical axis of the pipette. Five (17.9%) neurons with average location 145 (+/- 75) microns from the vertical axis exhibited both effects. Two (7.1%) neurons that exhibited significant increases in response amplitude to stimulus angles within 10 deg of the peak excitatory stimulus without changes in orientation selectivity or tuning were located less than 100 microns from the vertical axis. The effects on the orientation tuning of cells were restricted in all cases to within +/- 30 deg of the preferred stimulus orientation. This suggests that layer blocking affects cells with preferred stimulus orientations similar to those of the recorded neurons. Only cells located within 500 microns tangential to the vertical axis of the injection site exhibited these effects. These results suggest that cells within layers 5 and 6 provide organized, orientation-tuned inhibition that sharpens the orientation tuning of cells in the upper cortical layers within the same, or closely neighboring, cell columns.

Animals↗

GABAB-receptor-mediated inhibition reduces the orientation selectivity of the sustained response of striate cortical neurons in cats.

Blocking GABAA-receptor-mediated inhibition reduces the selectivity of striate cortical neurons for the orientation of a light bar primarily by reducing the selectivity of their onset transient (initial 200 ms) response. Blocking GABAB-receptor-mediated inhibition with phaclofen, however, is not reported to reduce the orientation selectivity of these neurons when it is measured with a light bar. We hypothesized that blocking GABAB-receptor-mediated inhibition would instead affect the orientation selectivity of cortical neurons by reducing the selectivity of their sustained response to a prolonged stimulus. To test this hypothesis, we stimulated 21 striate cortical neurons with drifting sine-wave gratings and measured their orientation selectivity before, during, and after iontophoretic injection of 2-hydroxy-saclofen (2-OH-S), a selective GABAB-receptor antagonist. 2-OH-S reduced the orientation selectivity of six of eight simple cells by an average of 28.8 (+/- 13.2) % and reduced the orientation selectivity of eight of 13 complex cells by an average of 32.3 (+/- 27.4) %. As predicted, 2-OH-S reduced the orientation selectivity of the neurons' sustained response, but did not reduce the orientation selectivity of their onset transient response. 2-OH-S also increased the length of spike "bursts" (two or more spikes with interspike intervals < or = 8 ms) and eliminated the orientation selectivity of these bursts for six cells. These results are the first demonstration of a functional role for GABAB receptors in visual cortex and support the hypothesis that two GABA-mediated inhibitory mechanisms, one fast and the other slow, operate within the striate cortex to shape the response properties of individual neurons.

Action Potentials↗

Contrast adaptation in striate cortical neurons of the nocturnal primate bush baby (Galago crassicaudatus).

It has been argued that in order for the visual system to detect edges accurately under a range of conditions, the visual system needs to adapt to the local contrast level to preserve sensitivity (Blakemore & Campbell, 1969). Cells in the primary visual cortex of cats adapt to stimuli with low to moderate contrast. Curiously, macaque monkey neurons in primary visual cortex (V1) do not show evidence for similar adaptation. To address the question of whether this differential sensitivity in contrast adaptation might be due to phylogenetic variation between cats and primates or to specializations for visual niche (e.g. nocturnal vs. diurnal), contrast adaptation to temporally and spatially optimized gratings was examined in 30 V1 cells of three nocturnal primate bush babies (Galago crassicaudatus). A second objective was to examine the relationship between the degree of contrast adaptation and cell classification or cell location relative to cortical layers or compartments [i.e. cytochrome-oxidase (CO) blobs and interblobs]. All cells were classified (simple vs. complex) and anatomically localized relative to cortical layers and cytochrome-oxidase (CO) blob and interblob compartments. Two independent measures of contrast adaptation were used. In the first test, contrast was sequentially increased from 3-56% and then decreased. The contrast required to maintain a half-maximum response amplitude in the 30 cells tested increased an average of 0.24 (+/- 0.12) log units during the sequential decrements in contrast. For the second test, four sets of five interleaved contrasts within +/- 1 octave of a central adapting contrast (10%, 14%, 20%, and 28%, respectively) were presented. The cells produced a mean adaptation index of 0.57 (+/- 0.47) which is very similar to that exhibited by cat cortical neurons (0.54 +/- 0.41). Interestingly, cells in interblobs showed a trend toward greater adaptation than did blob cells. Moreover, cells in the supragranular layers exhibited greater adaptation than cells in the infragranular layers. No significant differences in adaptation were found to correlate with other cell classification indices. Taken together, our results suggest that contrast adaptation may be more important for maintaining sensitivity in nocturnal species (primates or cats) than in diurnal species (macaque monkeys), and that in the nocturnal bush baby, cells in cortical layers and compartments may be differentially specialized for contrast adaptation.

Adaptation, Ocular↗

Differential contributions of magnocellular and parvocellular pathways to the contrast response of neurons in bush baby primary visual cortex (V1).

How neurons in the primary visual cortex (V1) of primates process parallel inputs from the magnocellular (M) and parvocellular (P) layers of the lateral geniculate nucleus (LGN) is not completely understood. To investigate whether signals from the two pathways are integrated in the cortex, we recorded contrast-response functions (CRFs) from 20 bush baby V1 neurons before, during, and after pharmacologically inactivating neural activity in either the contralateral LGN M or P layers. Inactivating the M layer reduced the responses of V1 neurons (n = 10) to all stimulus contrasts and significantly elevated (t = 8.15, P < 0.01) their average contrast threshold from 8.04 (+/- 4.1)% contrast to 22.46 (+/- 6.28)% contrast. M layer inactivation also significantly reduced (t = 4.06, P < 0.01) the average peak response amplitude. Inactivating the P layer did not elevate the average contrast threshold of V1 neurons (n = 10), but significantly reduced (t = 4.34, P < 0.01) their average peak response amplitude. These data demonstrate that input from the M pathway can account for the responses of V1 neurons to low stimulus contrasts and also contributes to responses to high stimulus contrasts. The P pathway appears to influence mainly the responses of V1 neurons to high stimulus contrasts. None of the cells in our sample, which included cells in all output layers of V1, appeared to receive input from only one pathway. These findings support the view that many V1 neurons integrate information about stimulus contrast carried by the LGN M and P pathways.

Animals↗

Are primate lateral geniculate nucleus (LGN) cells really sensitive to orientation or direction?

There is considerable controversy over the existence of orientation and direction sensitivity in lateral geniculate nucleus (LGN) neurons. Claims for the existence of these properties often were based upon data from cells tested well beyond their peak spatial frequencies. The goals of the present study were to examine the degree of orientation and direction sensitivity of LGN cells when tested at their peak spatial and temporal frequencies and to compare the tuning properties of these subcortical neurons with those of visual cortex. For this investigation, we used conventional extracellular recording to study orientation and direction sensitivities of owl monkey LGN cells by stimulating cells with drifting sinusoidal gratings at peak temporal frequencies, peak or higher spatial frequencies, and moderate contrast. A total of 110 LGN cells (32 koniocellular cells, 34 magnocellular cells, and 44 parvocellular cells) with eccentricities ranging from 2.6 deg to 27.5 deg were examined. Using the peak spatial and temporal frequencies for each cell, 41.8% of the LGN cells were found to be sensitive to orientation and 19.1% were direction sensitive. The degree of bias for orientation and direction did not vary with eccentricity or with cell class. Orientation sensitivity did, however, increase, and in some cases orientation preferences changed, at higher spatial frequencies. Increasing spatial frequency had no consistent effect on direction sensitivity. Compared to cortical cell orientation tuning, the prevalence and strength of LGN cell orientation and direction sensitivity are weak. Nevertheless, the high percentage of LGN cells sensitive to orientation even at peak spatial and temporal frequencies reinforces the view that subcortical biases could, in combination with activity-dependent cortical mechanisms and/or cortical inhibitory mechanisms, account for the much narrower orientation and direction tuning seen in visual cortex.

Animals↗

Modeling receptive-field structure of koniocellular, magnocellular, and parvocellular LGN cells in the owl monkey (Aotus trivigatus).

Most cells in the retina and lateral geniculate nucleus (LGN) of primates have a concentric center/surround receptive-field organization. Details of the relationship between center and surround often can be used to predict how cells respond to visual stimuli. Models of the receptive-field organization and center/surround relationships also are useful when comparing cell classes. In the present study, we used the difference-of-Gaussians (DOG) model to quantitatively examine and compare the receptive-field center/surround organization of koniocellular (K), magnocellular (M), and parvocellular (P) LGN cells of owl monkeys. We obtained estimates of receptive-field center size (r(c)) and center sensitivity (K(c)), and surround size (r(s)), and surround sensitivity (K(s)) from 62 K, M, and P LGN cells by fitting their spatial-frequency responses with a DOG function (Rodieck, 1965; Croner & Kaplan, 1995). The DOG function not only accounted for the responses of P and M cells, but also provided a good description of K-cell responses. We found that at matched eccentricities of less than 15 deg, K cells had the largest r(c) and r(s) among the three cell classes. K cells also had the lowest K(c) and K(s). Center and surround sizes tended to increase with retinal eccentricity for all three cell classes, but K cells showed a more variable pattern. There was an inverse relationship between sensitivity and size for both the receptive-field center and surround in all three cell classes. The surround/center volume ratio remained similar across cell classes. We conclude that K, M, and P LGN cell classes differ in the details of their receptive-field structure, but share common principles of center/surround organization.

Afferent Pathways↗

Nonlocal origin of response suppression from stimulation outside the classic receptive field in area 17 of the cat.

A stimulus located outside the classic receptive field (CRF) of a striate cortical neuron can markedly influence its behavior. To study this phenomenon, we recorded from two cortical sites, recorded and peripheral, with separate electrodes in cats anesthetized with Propofol and nitrous oxide. The receptive fields of each site were discrete (2-7.3 deg between centers). A control orientation tuning (OT) curve was measured for a single recorded cell with a drifting grating. The OT curve was then remeasured while stimulating simultaneously the cell's CRF as well as the peripheral site with a stimulus optimized for that location. For 22/60 cells, the peripheral stimulus suppressed the peak response and/or shifted the center of mass of the OT curve. For 19 of these 22 cells, we then reversibly blocked stimulus-driven activity at the peripheral site by iontophoretic application of GABA (0.5 M). For 6/19 cells, the response returned to control levels, implying that for these cells the inhibitory influence arose from the blocked site. The responses of nine cells remained reduced during inactivation of the peripheral site, suggesting that influence was generated outside the region of local block in area 17. This is consistent with earlier findings suggesting that modulatory influences can originate from higher cortical areas. Three cells had mixed results, suggesting multiple origins of influence. The response of each cell returned to suppressed levels after dissipation of the GABA and returned to baseline values when the peripheral stimulus was removed. These findings support a cortical model in which a cell's response is modulated by an inhibitory network originating from beyond the receptive field that supplants convergence of excitatory lateral geniculate neurons. The existence of cells that exhibit no change in peripherally inhibited responses during the GABA application suggests that peripheral influences may arise from outside area 17, presumably from other cortical areas (e.g. area 18).

Action Potentials↗

Temporal-frequency tuning of cross-orientation suppression in the cat striate cortex.

A sinusoidal mask grating oriented orthogonally to and superimposed onto an optimally oriented base grating reduces a cortical neuron's response amplitude. The spatial selectivity of cross-orientation suppression (XOR) has been described, so for this paper we investigated the temporal properties of XOR. We recorded from single striate cortical neurons (n = 72) in anesthetized and paralyzed cats. After quantifying the spatial and temporal characteristics of each cell's excitatory response to a base grating, we measured the temporal-frequency tuning of XOR by systematically varying the temporal frequency of a mask grating placed at a null orientation outside of the cell's excitatory orientation domain. The average preferred temporal frequency of the excitatory response of the neurons in our sample was 3.8 (+/- 1.5 S.D.) Hz. The average cutoff frequency for the sample was 16.3 (+/- 1.7) Hz. The average preferred temporal frequency (7.0 +/- 2.6 Hz) and cutoff frequency (20.4 +/- 6.9 Hz) of the XOR were significantly higher. The differences averaged 1.1 (+/- 0.6) octaves for the peaks and 0.3 (+/- 0.4) octaves for the cutoffs. The XOR mechanism's preference for high temporal frequencies suggests a possible extrastriate origin for the effect and could help explain the low-pass temporal-frequency response profile displayed by most striate cortical neurons.

Animals↗