Fixational eye movements in passive versus active sustained fixation tasks.

Human fixational eye movements are so small and precise that high-speed, accurate tools are needed to fully reveal their properties and functional roles. Where the fixated image lands on the retina and how it moves for different levels of visually demanding tasks is the subject of the current study. An Adaptive Optics Scanning Laser Ophthalmoscope (AOSLO) was used to image, track and present a variety of fixation targets (Maltese cross, disk, concentric circles, Vernier and tumbling-E letter) to healthy subjects. During these different passive (static) or active (discriminating) tasks under natural eye motion, the landing position of the target on the retina was tracked in space and time over the retinal image directly with high spatial (<1 arcmin) and temporal (960 Hz) resolution. We computed both the eye motion and the exact trajectory of the fixated target's motion over the retina. We confirmed that compared to passive tasks, active tasks elicited a partial inhibition of microsaccades, leading to longer drift periods compensated by larger corrective saccades. Consequently, the overall fixation stability during active tasks was on average 57% larger than during passive tasks. The preferred retinal locus of fixation was the same for each task and did not coincide with the location of the peak cone density.

The effects of fixational tremor on the retinal image.

The study of fixational eye motion has implications for the neural and computational underpinnings of vision. One component of fixational eye motion is tremor, a high-frequency oscillatory jitter reported to be anywhere from ∼11-60 arcseconds in amplitude. In order to isolate the effects of tremor on the retinal image directly and in the absence of optical blur, high-frequency, high-resolution eye traces were collected in six subjects from videos recorded with an adaptive optics scanning laser ophthalmoscope. Videos were acquired while subjects engaged in an active fixation task where they fixated on a tumbling E stimulus and reported changes in its orientation. Spectral analysis was conducted on periods of ocular drift, with all drifts being concatenated together after removal of saccades from the trace. The resultant amplitude spectra showed a slight deviation from the traditional 1/f nature of optical drift in the frequency range of 50-100 Hz, which is indicative of tremor. However, this deviation rarely exceeded 1 arcsecond and the consequent standard deviation of retinal image motion over the tremor band (50-100 Hz) was just over 5 arcseconds. Given such a small amplitude, it is unlikely tremor will contribute in any meaningful way to the visual percept.

Consequences of eye movements for spatial selectivity.

What determines spatial tuning in the visual system? Standard views rely on the assumption that spatial information is directly inherited from the relative position of photoreceptors and shaped by neuronal connectivity.1,2 However, human eyes are always in motion during fixation,3,4,5,6 so retinal neurons receive temporal modulations that depend on the interaction of the spatial structure of the stimulus with eye movements. It has long been hypothesized that these modulations might contribute to spatial encoding,7,8,9,10,11,12 a proposal supported by several recent observations.13,14,15,16 A fundamental, yet untested, consequence of this encoding strategy is that spatial tuning is not hard-wired in the visual system but critically depends on how the fixational motion of the eye shapes the temporal structure of the signals impinging onto the retina. Here we used high-resolution techniques for eye-tracking17 and gaze-contingent display control18 to quantitatively test this distinctive prediction. We examined how contrast sensitivity, a hallmark of spatial vision, is influenced by fixational motion, both during normal active fixation and when the spatiotemporal stimulus on the retina is altered to mimic changes in fixational control. We showed that visual sensitivity closely follows the strength of the luminance modulations delivered within a narrow temporal bandwidth, so changes in fixational motion have opposite visual effects at low and high spatial frequencies. By identifying a key role for oculomotor activity in spatial selectivity, these findings have important implications for the perceptual consequences of abnormal eye movements, the sources of perceptual variability, and the function of oculomotor control.

Microsaccades during reading.

Recent research has shown that microsaccades contribute to high acuity vision. However, little is known about whether microsaccades also play a role in daily activities, such as reading, that do not involve stimuli at the limit of spatial resolution. While the functions of larger saccades in reading have been extensively examined, microsaccades are commonly regarded as oculomotor noise in this context. We used high-resolution eyetracking and precise gaze localization to investigate fine oculomotor behavior during reading. Our findings show that microsaccade characteristics differ from those measured during sustained fixation: microsaccades are larger in size and primarily leftwards during reading, i.e. they move the line of sight backward on the text. Analysis of how microsaccades shift gaze relative to the text suggests that these movements serve two important functions: (1) a corrective function, by moving the gaze regressively within longer words when the preceding saccade lands too far toward the end of these words, and (2) an exploratory function, by shifting the gaze on adjacent words to gain additional information before the execution of the next saccade. Thus, microsaccades may benefit reading by enhancing the visibility of nearby words. This study highlights the importance of examining fine oculomotor behavior in reading, and calls for further research to investigate the possible roles of microsaccades in reading difficulties.