Biomedical subjects

Michael J Hawken

Publications and source records attributed to Michael J Hawken.

12 recordsLinked to original sources

PubMed2005

Effects of contrast on smooth pursuit eye movements.

It is well known that moving stimuli can appear to move more slowly when contrast is reduced (P. Thompson, 1982). Here we address the question whether changes in stimulus contrast also affect smooth pursuit eye movements. Subjects were asked to smoothly track a moving Gabor patch. Targets varied in velocity (1, 8, and 15 deg/s), spatial frequency (0.1, 1, 4, and 8 c/deg), and contrast, ranging from just below individual thresholds to maximum contrast. Results show that smooth pursuit eye velocity gain rose significantly with increasing contrast. Below a contrast level of two to three times threshold, pursuit gain, acceleration, latency, and positional accuracy were severely impaired. Therefore, the smooth pursuit motor response shows the same kind of slowing at low contrast that was demonstrated in previous studies on perception.

Contrast Sensitivity↗

PubMed2005

Functional maturation of the macaque's lateral geniculate nucleus.

Vision in infant primates is poor, but it is not known which structures in the eye or brain set the main limits to its development. We studied the visual response properties of 348 neurons recorded in the lateral geniculate nucleus (LGN) of macaque monkeys aged 1 week to adult. We measured spatial and temporal frequency tuning curves and contrast responses with drifting achromatic sinusoidal gratings. Even in animals as young as 1 week, the main visual response properties of neurons in the magnocellular (M) and parvocellular (P) divisions of the LGN were qualitatively normal, including the spatial organization of receptive fields and the characteristic response properties that differentiate M- and P-cells. At 1 and 4 weeks, spatial and temporal resolution were less than one-half of adult values, whereas contrast gain and peak response rates for optimal stimuli were about two-thirds of adult values. Adult levels were reached by 24 weeks. Analysis of correlations between S-potentials representing retinal inputs and LGN cells suggested that the LGN follows retinal input as faithfully in infants as in adults, implicating retinal development as the main driving force in LGN development. Comparisons with previously published psychophysical data and ideal observer models suggest that the relatively modest changes in LGN responses during maturation impose no significant limits on visual performance. In contrast to previous studies, we conclude that these limits are set by neural development in the visual cortex, not in or peripheral to the LGN.

Age Factors↗

PubMed2005

Effect of stimulus size on the dynamics of orientation selectivity in Macaque V1.

Previous research has established that orientation selectivity depends to a great extent on suppressive mechanisms in the visual cortex. In this study, we investigated the spatial organization and the time-course of these mechanisms. Stimuli were presented in circular windows of "optimal" and "large" radii. The two stimulus sizes were chosen based on an area-response function measured with drifting gratings at high contrast. The "optimal" size was defined as the smallest radius that elicited the peak response (average value of 0.45 degrees ), whereas "large" was defined as two to five times the optimal size. We found that the peak amplitude of tuned enhancement and untuned suppression varied <10% on average with stimulus radius, indicating that they are mainly concentrated in the classical receptive field. However, tuned suppression--in those cells that showed it--was significantly stronger with large stimuli, indicating that this component has a contribution from beyond the classical receptive field. These results imply that spatial context (in large stimuli) enhances orientation selectivity by increasing tuned suppression. We also characterized the time evolution of enhancement, of untuned suppression, and of tuned suppression. The time-course of tuned suppression was markedly slower in time-to-peak and longer in its persistence than untuned suppression. Therefore tuned suppression is likely to be generated by long-range recurrent connections or cortico-cortical feedback, whereas untuned suppression is mainly generated locally in V1.

Animals↗

PubMed2004

Entrainment to video displays in primary visual cortex of macaque and humans.

Cathode ray tubes (CRTs) display images refreshed at high frequency, and the temporal waveform of each pixel is a luminance impulse only a few milliseconds long. Although humans are perceptually oblivious to this flicker, we show in V1 in macaque monkeys and in humans that extracellularly recorded action potentials (spikes) and visual-evoked potentials (VEPs) align with the video impulses, particularly when high-contrast stimuli are viewed. Of 91 single units analyzed in macaque with a 60 Hz video refresh, 29 cells (32%) significantly locked their firing to a uniform luminance display, but their number increased to 75 (82%) when high-contrast stimuli were shown. Of 92 cells exposed to a 100 Hz refresh, 21 (23%) significantly phase locked to high-contrast stimuli. Phase locking occurred in both input and output layers of V1 for simple and complex cells, regardless of preferred temporal frequency. VEPs recorded in humans showed significant phase locking to the video refresh in all seven observers. Like the monkey neurons, human VEPs more typically phase locked to stimuli containing spatial contrast than to spatially uniform stimuli. Phase locking decreased when the refresh rate was increased. Thus in humans and macaques phase locking to the high strobe frequency of a CRT is enhanced by a salient spatial pattern, although the perceptual impact is uncertain. We note that a billion people worldwide manage to watch TV without obvious distortion of their visual perception despite extraordinary phase locking of their V1s to a 50 or 60 Hz signal.

Action Potentials↗

PubMed2004

Correlation of local and global orientation and spatial frequency tuning in macaque V1.

Visual cortical neurones display a variety of visual properties. Among those that emerge in the primary visual cortex V1 are sharpening of selectivity for spatial frequency and for orientation. The selectivity for these stimulus attributes can be measured around the peak of the tuning function, usually as bandwidth. Other selectivity measures take into account the response across a broader range of stimulus values. An example of such a global measure is the circular variance of orientation tuning. Here we introduce a similar measure in the spatial frequency domain that takes into account the shape of the tuning curve at frequencies lower than the peak, called the low-spatial frequency variance. Our recent studies with dynamic stimuli suggest that the selectivity for spatial frequency and orientation is strongly correlated with the degree of suppression at low spatial frequencies and off-axis orientations. Here we extend the study of the global tuning to stimulus conditions that measure the response of cells to the presentation of drifting sinusoidal grating stimuli for periods of a few seconds. We find that under such steady-state stimulus conditions there is a strong correlation between the global selectivity measures, orientation circular variance and low spatial frequency variance. Consistent with previous studies, there is a weaker correlation between the local tuning measures, orientation and spatial frequency bandwidth. These results support the idea that there are multiple factors that contribute to tuning and that suppression observed in dynamic experiments is also likely to contribute to the global selectivity for steady-state stimuli.

Animals↗

PubMed2004

Cone inputs in macaque primary visual cortex.

To understand the role of primary visual cortex (V1) in color vision, we measured directly the input from the 3 cone types in macaque V1 neurons. Cells were classified as luminance-preferring, color-luminance, or color-preferring from the ratio of the peak amplitudes of spatial frequency responses to red/green equiluminant and to black/white (luminance) grating patterns, respectively. In this study we used L-, M-, and S-cone-isolating gratings to measure spatial frequency response functions for each cone type separately. From peak responses to cone-isolating stimuli we estimated relative cone weights and whether cone inputs were the same or opposite sign. For most V1 cells the relative S-cone weight was <0.1. All color-preferring cells were cone opponent and their L/M cone weight ratio was clustered around a value of -1, which is roughly equal and opposite L and M cone signals. Almost all cells (88%) classified as luminance cells were cone nonopponent, with a broad distribution of cone weights. Most cells (73%) classified as color-luminance cells were cone opponent. This result supports our conclusion that V1 color-luminance cells are double-opponent. Such neurons are more sensitive to color boundaries than to areas of color and thereby could play an important role in color perception. The color-luminance population had a broad distribution of L/M cone weight ratios, implying a broad distribution of preferred colors for the double-opponent cells.

Action Potentials↗

PubMed2003

A comparison of pursuit eye movement and perceptual performance in speed discrimination.

Currently there is considerable debate as to the nature of the pathways that are responsible for the perception and motor performance. We have studied the relationship between perceived speed, which is the experiential representation of a moving stimulus, and the speed of smooth pursuit eye movements, the motor action. We determined psychophysical thresholds for detecting small perturbations in the speed of moving patterns, and then by an ideal observer analysis computed analogous "oculometric" thresholds from the eye movement traces elicited by the same stimuli on the same trials. Our results confirm those of previous studies that show a remarkable agreement between perceptual judgments for speed discrimination and the fine gradations in eye movement speed. We analyzed the initial pursuit period of long duration (1000 ms) and short (200 ms) duration perturbations. When we compared the errors for perception and pursuit on a trial-by-trial basis there was no correlation between perceptual errors and eye movement errors. The observation that both oculometric and psychometric performance were similar, with Weber fractions in the normal range, but that there is no correlation in the errors suggests that the motor system and perception share the same constraints in their analysis of motion signals, but act independently and have different noise sources. We simulated noise in two models of perceptual and eye movement performance. In the first model we postulate an initial common source for the perceptual and eye movement signals. In that case about ten times the observed noise is required to produce no correlation in trial-by-trial performance. In the second model we postulate that the perceptual signal is a combination of a reafferent eye velocity signal plus the perturbation signal while the pursuit signal is derived from the oculomotor plant plus the perturbation signal. In this model about three times the noise level in the independent signals will mask any correlation due to the common perturbation signal.

Humans↗

PubMed2003

Ideal observer analysis of the development of spatial contrast sensitivity in macaque monkeys.

To explore the factors limiting the development of visual sensitivity, we constructed an ideal observer model for the infant macaque visual system. We made measurements of retinal morphology in infant and adult macaque monkeys, and used the data in combination with published optical data to formulate the model. We compared the ideal observer's ability to detect low-contrast gratings presented either in isolation or in spatiotemporal noise with behavioral data obtained under matched conditions. The ideal observer showed some improvement in visual performance up to the age of 4 weeks, but little change thereafter. Behavioral data show extensive changes over the ages 5-50 wk, after the ideal observer's performance has become asymptotic. We conclude that the development of visual sensitivity in infant monkeys is not limited by changes in the front-end factors captured by the ideal observer model, at least after the age of 5 weeks. Using noise masking, we also estimated the variability of neural processing in comparison with the photon noise-limited ideal. We found that both additive and multiplicative components of this variability are elevated in infant monkeys, and improve (though not to ideal levels) during development. We believe that these changes all reflect maturation of visual processing in cortical circuits, and that no aspect of visual performance in the regime we studied is limited by the properties of the retina and photoreceptors, either in infant or in adult animals.

Animals↗

PubMed2003

Dynamics of orientation tuning in macaque V1: the role of global and tuned suppression.

The temporal development of neural selectivity to physical attributes of a visual stimulus, such as its orientation and spatial frequency, can provide important clues about mechanisms of cortical tuning. We measured the dynamics of orientation tuning in macaque primary visual cortex (V1) and found several dynamical features in the data: changes in global enhancement and suppression, narrowing of orientation bandwidth, small but significant shifts in preferred orientation, and "Mexican-hat" tuning curves. The dynamics data were analyzed with a model that sums two fixed, tuned components (enhancement and suppression) and one global (untuned) component. The analysis suggests that there is early global enhancement followed by global and tuned suppression. Tuned suppression accounts for the dynamical reduction of orientation bandwidth and for the generation of Mexican-hat tuning profiles. Our findings imply that global and tuned suppression are important factors that determine the selectivity and dynamics of V1 responses to orientation.

Animals↗

PubMed2002

Orientation selectivity in macaque V1: diversity and laminar dependence.

We studied the steady-state orientation selectivity of single neurons in macaque primary visual cortex (V1). To analyze the data, two measures of orientation tuning selectivity, circular variance and orientation bandwidth, were computed from the tuning curves. Circular variance is a global measure of the shape of the tuning curve, whereas orientation bandwidth is a local measure of the sharpness of the tuning curve around its peak. Circular variance in V1 was distributed broadly, indicating a great diversity of orientation selectivity. This diversity was also reflected in the individual cortical layers. However, there was a tendency for neurons with high circular variance, meaning low selectivity for orientation, to be concentrated in layers 4C, 3B, and 5. The relative variation of orientation bandwidth across the cortical layers was less than for circular variance, but it showed a similar laminar dependence. Neurons with large orientation bandwidth were found predominantly in layers 4C and 3B. There was a weak correlation between orientation selectivity and the level of spontaneous activity of the neurons. We also assigned a response modulation ratio for each cell, which is a measure of the linearity of spatial summation. Cells with low modulation ratios tended to have higher circular variance and bandwidth than those with high modulation ratios. These findings suggest a revision to the classical view that nonoriented receptive fields are principally found in layer 4C and the cytochrome oxidase-rich blobs in layer 2/3. Instead, a broad distribution of tuning selectivity is found in all cortical layers, and neurons that are weakly tuned for orientation are ubiquitous in V1 cortex.

Action Potentials↗

PubMed2002

Contrast-dependent changes in spatial frequency tuning of macaque V1 neurons: effects of a changing receptive field size.

Previous studies on single neurons in primary visual cortex have reported that selectivity for orientation and spatial frequency tuning do not change with stimulus contrast. The prevailing hypothesis is that contrast scales the response magnitude but does not differentially affect particular stimuli. Models where responses are normalized over contrast to maintain constant tuning for parameters such as orientation and spatial frequency have been proposed to explain these results. However, our results indicate that a fundamental property of receptive field organization, spatial summation, is not contrast invariant. We examined the spatial frequency tuning of cells that show contrast-dependent changes in spatial summation and have found that spatial frequency selectivity also depends on stimulus contrast. These results indicate that contrast changes in the spatial frequency tuning curves result from spatial reorganization of the receptive field.

Animals↗

PubMed2002

Receptive field structure of neurons in monkey primary visual cortex revealed by stimulation with natural image sequences.

Probing the visual system with the ensemble of signals that occur in the natural environment may reveal aspects of processing that are not evident in the neural responses to artificial stimulus sets, such as conventional bars and sinusoidal gratings. However, unsolved is the question of how to use complex natural stimulation, many aspects of which the experimenter cannot completely specify, to study neural processing. Here a method is presented to investigate the structure of a neuron's receptive field based on its response to movie clips and other stimulus ensembles. As a particular case, the technique provides an estimate of the conventional first-order receptive field of a neuron, similar to what can be obtained with other reverse-correlation schemes. This is demonstrated experimentally and with computer simulations. Our analysis also revealed that the receptive fields of both simple and complex cells had regions where image boundaries, independent of their contrast sign, would enhance or suppress the cell's response. In some cases, these signals were tuned for the orientation of the boundary. This demonstrates for the first time that it might be feasible to investigate the receptive field structure of visual neurons from their responses to natural image sequences.

Animals↗