Attentional shift

Some of the first research into the neurology behind attention shifts came from examining brain damaged patients. First, Posner et al., studied persons affected by progressive supranuclear palsy, a condition wherein it is difficult to exert eye movements voluntarily, particularly vertical movements. Patients were found to have damage present in the mid-brain area and associated cortical areas. Although patients were not able to move their eyes, they were still able to shift attention covertly. However, there was a slowing of the process of shifting attention in these patients, suggesting that the mid-brain and cortical areas must be associated with covert attention shifts. Additionally, previous research has shown support for covert attention shifts being associated with activity in the parietal lobe. On the other hand, research seems to indicate differences in brain areas activated for overt attention shifts, as compared to covert shifts. Previous evidence has shown that the superior colliculus is associated with eye movements, or overt attention shifts.[16] Additionally, the medial cerebellum has shown activation only during eye movements.[17]

Although, after reviewing Posner's research, it may seem logical to conclude that covert and overt attention shifts utilize different neural mechanisms, other more recent studies have shown more overlap than not. Multiple studies have shown activity evident in the frontal cortex, concentrating in the precentral sulcus, the parietal cortex, specifically in the intraparietal sulcus, and in the lateral occipital cortex for both overt and covert attention shifts.[18] This is in support of the premotor theory of attention. While these studies may agree on the areas, they are not always in agreement on whether an overt or covert attentional shift causes more activation. Utilizing functional magnetic resonance imaging (fMRI) technology, Corbetta et al., found that overt and covert attention shift tasks showed activation within the same areas, namely, the frontal, parietal and temporal lobes. Additionally, this study reported that covert shifts of attention showed greater activity levels than in the overt attention condition. However, it is important to note that different tasks were used for the covert versus the overt condition. One task involved a probe being flashed to the subject's fovea, while another task showed the probe in the participant's peripheral vision, making it questionable whether these results can be directly compared.[17] Nobre et al. also sought to determine whether covert and overt attention shifts revealed activation in the same brain areas. Once again fMRI technology was utilized, as well as, two separate tasks, one for covert attention and one for overt attention. Results showed overlap in activated areas for overt and covert attention shifts, mainly in the parietal and frontal lobes. However, one area was shown to be specific to covert attention, which was the right dorsolateral cortex; typically associated with voluntary attention shifts and working memory. One should question whether this additional activation has to do with the selected task for the covert condition, or rather if it is specific to a covert shift of attention.[19]

Beauchamp et al. more recently attempted to reproduce these same results by performing a study utilizing the same task for both conditions, as well as across multiple shift rates. Results were in agreement that covert and overt attentional shifts engage the same neural mechanisms. However, this study differed in that overt shifts of attention showed greater activation in these neural areas, and this occurred even at multiple shift rates. Once again, the neural regions implicated in this study included the intraparietal sulcus, the precentral sulcus, and the lateral occipital cortex. This larger activation evident with overt attention shifts was attributed to the added involvement of eye movements.[18]

Corbetta and Shulman, who are proponents of the belief that separate neural systems exist for endogenous and exogenous control, conducted a meta-analysis of multiple studies showing brain activation due to either of the two attentional processes. Specifically, the dorsal posterior parietal and frontal cortex region are mainly implicated with voluntary attention, while activity is transiently shown in the occipital region. The endogenous mechanisms are thought to integrate previous knowledge, expectations and goals to voluntarily decide where to shift attention. On the other hand, neural areas involved in reflexive attention are believed to have the purpose of focusing attention on events or objects that stand out in the environment. The temporoparietal cortex and ventral frontal cortex region, particularly in the right brain hemisphere, have shown involvement with reflexive attention.[21] One kind of visual inputs stands out for the primary visual cortex (V1) but not for visual awareness or for other cortical areas,[22] they are distinctive in term of whether the left or right eye receives the inputs, e.g., an apple shown to the left eye among many other apples of the same appearance shown to the right eye. Nevertheless, such inputs, e.g., the left-eye apple, can also strongly capture attention overly and covertly (even overriding attentional guidance by endogenous goals),[23][24] implicating V1 for exogenous attentional shifts according to V1 Saliency Hypothesis.[25] Even though separate regions are thought to be in existence for these two attentional processes, the question still remains on whether these regions interact with one another, indicating more research on this point is still needed.[9]

There appears to be agreement that multiple areas of the brain are involved in shifts of attention, however research is not quite as conclusive regarding the amount of overlap evident with voluntary versus reflexive attention. Rosen et al.'s study found a fair amount of overlap between endogenous and exogenous shifts of attention. Both conditions showed activation in the dorsal and parietal premotor areas. However, the voluntary condition also showed activation in the right dorsolateral prefrontal cortex, which did not appear in the reflexive condition. As this area has been shown to be associated with working memory, it may indicate that working memory is engaged voluntarily. The subcortical global pallidus region was also activated only in the voluntary condition. Additionally, the activation shown in the temporoparietal junction [TPJ] was slightly different in both conditions, with the endogenous condition showing more spreading to the lateral, anterior and superior regions. Although these differences did exist, overall there was a lot of overlap demonstrated for voluntary and reflexive shifts of attention. Specifically both showed activations in the dorsal premotor region, the frontal eye field area, and the superior parietal cortex (SPC), although, the SPC exhibited greater activation in the endogenous condition.[26]

Attention can be guided by top-down processing or via bottom up processing. Posner's model of attention includes a posterior attentional system involved in the disengagement of stimuli via the parietal cortex, the shifting of attention via the superior colliculus and the engagement of a new target via the pulvinar. The anterior attentional system is involved in detecting salient stimuli and preparing motor responses.