Brian J. White

Color-Related Signals in the Primate Superior Colliculus

Color is important for segmenting objects from backgrounds, which can in turn facilitate visual search in complex scenes. However, brain areas involved in orienting the eyes toward colored stimuli in our environment are not believed to have access to color information. Here, we show that neurons in the intermediate layers of the monkey superior colliculus (SC), a critical structure for the production of saccadic eye movements, can respond to isoluminant color stimuli with the same magnitude as a maximum contrast luminance stimulus. In contrast, neurons from the superficial SC layers showed little color-related activity. Crucially, visual onset latencies were 30–35 ms longer for color, implying that luminance and chrominance information reach the SC through distinct pathways and that the observed color-related activity is not the result of residual luminance signals. Furthermore, these differences in visual onset latency translated directly into differences in saccadic reaction time. The results demonstrate that the saccadic system can signal the presence of chromatic stimuli only one stage from the brainstem premotor circuitry that drives the eyes.

Microstimulation of the Monkey Superior Colliculus Induces Pupil Dilation Without Evoking Saccades

The orienting reflex is initiated by a salient stimulus and facilitates quick, appropriate action. It involves a rapid shift of the eyes, head, and attention and other physiological responses such as changes in heart rate and transient pupil dilation. The SC is a critical structure in the midbrain that selects incoming stimuli based on saliency and relevance to coordinate orienting behaviors, particularly gaze shifts, but its causal role in pupil dilation remains poorly understood in mammals. Here, we examined the role of the primate SC in the control of pupil dynamics. While requiring monkeys to keep their gaze fixed, we delivered weak electrical microstimulation to the SC, so that saccadic eye movements were not evoked. Pupil size increased transiently after microstimulation of the intermediate SC layers (SCi) and the size of evoked pupil dilation was larger on a dim versus bright background. In contrast, microstimulation of the superficial SC layers did not cause pupil dilation. Thus, the SCi is directly involved not only in shifts of gaze and attention, but also in pupil dilation as part of the orienting reflex, and the function of pupil dilation may be related to increasing visual sensitivity. The shared neural mechanisms suggest that pupil dilation may be associated with covert attention.

Linking visual response properties in the superior colliculus to saccade behavior

Here we examined the influence of the visual response in the superior colliculus (SC) (an oculomotor control structure integrating sensory, motor and cognitive signals) on the development of the motor command that drives saccadic eye movements in monkeys. We varied stimulus luminance to alter the timing and magnitude of visual responses in the SC and examined how these changes correlated with resulting saccade behavior. Increasing target luminance resulted in multiple modulations of the visual response, including increased magnitude and decreased response onset latency. These signal modulations correlated strongly with changes in saccade latency and metrics, indicating that these signal properties carry through to the neural computations that determine when, where and how fast the eyes will move. Thus, components of the earliest part of the visual response in the SC provide important building blocks for the neural basis of the sensory–motor transformation, highlighting a critical link between the properties of the visual response and saccade behavior.

Separate Visual Signals for Saccade Initiation during Target Selection in the Primate Superior Colliculus

The primary function of the superior colliculus (SC) is to orient the visual system toward behaviorally relevant stimuli defined by features such as color. However, a longstanding view has held that visual activity in the SC arises exclusively from achromatic pathways. Recently, we reported evidence that the primate SC is highly sensitive to signals originating from chromatic pathways, but these signals are delayed relative to luminance signals (White et al., 2009). Here, we describe a functional consequence of this difference in visual arrival time on the processes leading to target selection and saccade initiation. Two rhesus monkeys performed a simple color-singleton selection task in which stimuli carried a chromatic component only (target and distractors were isoluminant with the background, but differed in chromaticity) or a combined chromatic–achromatic component (36% luminance contrast added equally to all stimuli). Although visual responses were delayed in the chromatic-only relative to the combined chromatic–achromatic condition, SC neurons discriminated the target from distractors at approximately the same time provided stimulus chromaticity was held constant. However, saccades were triggered sooner, and with more errors, with the chromatic–achromatic condition, suggesting that luminance signals associated with these stimuli increased the probability of triggering a saccade before the target color was adequately discriminated. These results suggest that separate mechanisms may independently influence the saccadic command in the SC, one linked to the arrival time of pertinent visual signals, and another linked to the output of the visual selection process.

Interaction between visual-and goal-related neuronal signals on the trajectories of saccadic eye movements

During natural viewing, the trajectories of saccadic eye movements often deviate dramatically from a straight-line path between objects. In human studies, saccades have been shown to deviate toward or away from salient visual distractors depending on visual- and goal-related parameters, but the neurophysiological basis for this is not well understood. Some studies suggest that deviation toward is associated with competition between simultaneously active sites within the intermediate layers of the superior colliculus (SC), a midbrain structure that integrates sensory and goal-related signals for the production of saccades. In contrast, deviation away is hypothesized to reflect a higher-level process, whereby the neural site associated with the distractor isactively suppressed via a form of endogenous, top–down inhibition. We tested this hypothesis by measuring presaccadic distractor-evoked activation of SC visuomotor neurons while monkeys performed a simple task configured specifically to induce a high degree of saccades that deviate away. In the SC, cognitive processes such as top–down expectation are represented as variation in the sustained, low-frequency presaccadic discharge. We reasoned that any inhibition at the distractor-related locus associated with saccade deviation should affect the excitability of the neuron, thereby affecting the discharge rate. We found that, although the task produced robust deviation away, there was no evidence of a relationship between saccade deviation and distractor-evoked activation outside a short perisaccadic window that began no earlier than 22 msec before saccade onset. This indicates that deviation away is not adequately explained by a form of sustained, top–down inhibition at the distractor-related locus in the SC. The results are discussed in relation to the primary sources of inhibition associated with saccadic control.

Zero ripple single stage AC-DC LED driver with unity power factor

The single stage LED Driver can achieve relatively high efficiency, however the low frequency ripple current is too significant if high power factor has been achieved. With two stages AC-DC LED Driver structure, we can achieve high power factor and tight current regulation at the same time. However, the drawback of the two stage structure is relatively low efficiency and high component cost. In this paper, an innovative single stage LED Driver with ripple cancellation technology has been proposed. We can achieve almost as high efficiency and low component cost as single stage LED Driver while maintaining comparable performance to the two stages LED Driver. Our experimental prototype can achieve 1.5mA Pk-Pk 120Hz ripple current, 0.99 power factor and 85.5% efficiency for a universal AC input, 35W (50V-0.7A) output application.

The spatio-temporal tuning of the mechanisms in the control of saccadic eye movements.

We compared the spatio-temporal tuning of perception to the mechanisms that drive saccadic eye movements. Detection thresholds were measured for Gabor-targets presented left or right of fixation (4 or 8deg eccentricity), at one of four spatial frequencies (1, 2, 4 or 8cpd) oscillating at one of three temporal frequencies (1, 8 or 16Hz). We then measured saccade latency to each target presented at various multiples of detection threshold. Consistent with previous research, latency decreased as a function of contrast. However, at equal detection performance, we found no systematic difference in saccadic latency and no difference in average oculometric performance (% correct saccade direction) across the different target spatio-temporal frequencies. Furthermore, position error remained fairly constant across all conditions. The results are consistent with the idea that the spatio-temporal signals used for perception are the same as those used by the mechanisms driving saccadic eye movements.

Superior colliculus neurons encode a visual saliency map during free viewing of natural dynamic video

Models of visual attention postulate the existence of a saliency map whose function is to guide attention and gaze to the most conspicuous regions in a visual scene. Although cortical representations of saliency have been reported, there is mounting evidence for a subcortical saliency mechanism, which pre-dates the evolution of neocortex. Here, we conduct a strong test of the saliency hypothesis by comparing the output of a well-established computational saliency model with the activation of neurons in the primate superior colliculus (SC), a midbrain structure associated with attention and gaze, while monkeys watched video of natural scenes. We find that the activity of SC superficial visual-layer neurons (SCs), specifically, is well-predicted by the model. This saliency representation is unlikely to be inherited from fronto-parietal cortices, which do not project to SCs, but may be computed in SCs and relayed to other areas via tectothalamic pathways.

Saccadic facilitation in natural backgrounds.

In visual systems with a fovea, only a small portion of the visual field can be analyzed with high accuracy. Saccadic eye movements shift that center of gaze around several times a second. Saccades have been characterized in great detail and depend critically on a number of visual properties of the stimuli. However, typical experiments have used bright spots on dark backgrounds, while our natural environment has a highly characteristic rich spatial structure. Here we show that the saccadic system, unlike the perceptual system, is able to compensate for the masking caused by structured backgrounds. Consequently, saccadic latencies in the context of natural backgrounds are much faster than unstructured backgrounds at equal levels of visibility. The results suggest that whenever a structured background acts to mask the visibility of the saccade target, it simultaneously preactivates saccadic circuitry and thus ensures a fast reaction to potentially critical stimuli that are difficult to detect in our environment.

Superior colliculus encodes visual saliency before the primary visual cortex

Significance Theories of visual attention postulate the existence of a saliency map that guides attention/gaze toward the most visually conspicuous stimuli in complex scenes. This study compared saliency coding in the two dominant visual gateways: the primary visual cortex (V1) and the evolutionarily older visual system that exists in the midbrain superior colliculus. Our results show that neurons in the superficial visual layers of the superior colliculus (SCs) encoded saliency earlier and more robustly than V1 neurons. This was surprising, because the dominant input to the SCs arises from V1. This result is in line with models that place a feature processing stage (V1) before the feature-agnostic saliency map in SCs. Models of visual attention postulate the existence of a bottom-up saliency map that is formed early in the visual processing stream. Although studies have reported evidence of a saliency map in various cortical brain areas, determining the contribution of phylogenetically older pathways is crucial to understanding its origin. Here, we compared saliency coding from neurons in two early gateways into the visual system: the primary visual cortex (V1) and the evolutionarily older superior colliculus (SC). We found that, while the response latency to visual stimulus onset was earlier for V1 neurons than superior colliculus superficial visual-layer neurons (SCs), the saliency representation emerged earlier in SCs than in V1. Because the dominant input to the SCs arises from V1, these relative timings are consistent with the hypothesis that SCs neurons pool the inputs from multiple V1 neurons to form a feature-agnostic saliency map, which may then be relayed to other brain areas.