Recent evidence suggests that microsaccades are causally linked to the attention-related modulation of neurons—specifically, that microsaccades toward the attended location are required for the subsequent changes in firing rate. These findings have raised questions about whether attention-related modulation is due to different states of attention as traditionally assumed or might instead be a secondary effect of microsaccades. Here, in two rhesus macaques, we tested the relationship between microsaccades and attention-related modulation in the superior colliculus (SC), a brain structure crucial for allocating attention. We found that attention-related modulation emerged even in the absence of microsaccades, was already present prior to microsaccades toward the cued stimulus, and persisted through the suppression of activity that accompanied all microsaccades. Nonetheless, consistent with previous findings, we also found significant attention-related modulation when microsaccades were directed toward, rather than away from, the cued location. Thus, despite the clear links between microsaccades and attention, microsaccades are not necessary for attention-related modulation, at least not in the SC. They do, however, provide an additional marker for the state of attention, especially at times when attention is shifting from one location to another.

Even during fixation, our eyes are in constant motion. For example, microsaccades are small (typically <1) eye movements that occur 1~3 times/second. Despite their tiny and transient nature, our percept of visual space is compressed before microsaccades (Hafed ZM, Lovejoy LP, Krauzlis RJ. 37: 1169-1181, 2013). As visual space and time are interconnected at both the physical and physiological levels, we asked whether microsaccades also affect the temporal aspects of visual perception. Here we demonstrate that the perceived interval between transient visual stimuli was compressed if accompanied by microsaccades. This temporal compression extended approximately ±200 ms from microsaccade occurrence, and depending on their particular pattern, multiple microsaccades further enhanced or counteracted this temporal compression. The compression of time surrounding microsaccades resembles that associated with more voluntary macrosaccades (Morrone MC, Ross J, Burr D. 8: 950-954, 2005). Our results suggest common neural processes underlying both saccade and microsaccade misperceptions, mediated, likely, through extraretinal mechanisms. Here we show that humans perceive the duration of visual events as compressed if they are accompanied by microsaccades. Despite the tiny and transient nature of microsaccades, time compression extended more than ±200 ms from their occurrence. Moreover, the number, pattern, and temporal coincidence of microsaccades relative to visual events all contribute to this time misperception. Our results reveal a detailed picture of how our visual time percepts are altered by microsaccades.

We define attention as "the set of evolved brain processes that leads to adaptive and effective behavioral selection." Our emphasis is on understanding the biological and neural mechanisms that make the behavioral properties of attention possible. Although much has been learned about the functional operation of attention by postulating and testing different aspects of attention, our view is that the distinctions most frequently relied upon are much less useful for identifying the detailed biological mechanisms and brain circuits. Instead, we adopt an evolutionary perspective that, while speculative, generates a different set of guiding principles for understanding the form and function of attention.We then provide a thought experiment, introducing a device that we intend to serve as an intuition pump for thinking about how the brain processes for attention might be organized, and that illustrates the features of the biological processes that might ultimately answer the question.

Two spatial reference systems, i.e., the observer-centered (egocentric) and object-centered (allocentric) references, are most commonly used to locate the position of the external objects in space. Although we sense the world as a unified entity, visual processing is asymmetric between upper and lower visual fields (VFs). For example, the goal-directed reaching responses are more efficient in the lower VF. Such asymmetry suggests that the visual space might be composed of different realms regarding perception and action. Since the peripersonal realm includes the space that one can reach, mostly in the lower VF, it is highly likely that the peripersonal realm might mainly be represented in the egocentric reference for visuomotor operation. In contrast, the extrapersonal realm takes place away from the observer and is mostly observed in the upper VF, which is presumably represented in the allocentric reference for orientation in topographically defined space. This theory, however, has not been thoroughly tested experimentally. In the present study, we assessed the contributions of the egocentric and allocentric reference systems on visual discrimination in the upper and lower VFs through measuring the manual reaction times (RTs) of human subjects. We found that: (a) the influence of a target's egocentric location on visual discrimination was stronger in the lower VF; and (b) the influence of a target's allocentric location on visual discrimination was stronger in the upper VF. These results support the hypothesis that the upper and lower VFs are primarily represented in the allocentric and egocentric references, respectively.