Attention has a central function in the construction of every visual experience. This review considers the contributions of functional neuroimaging to our understanding of what visual attention is and how it works. It addresses four principle questions:
What is the locus of attentional selection? The debate concerning early versus late selection is evaluated in light of recent findings. New imaging evidence indicates that attention affects neural activity not only in extrastriate cortex, but also at the first stage of cortical information processing in primary visual cortex.
What exactly gets selected by attention? Behavioural data show that selection can operate at the level of spatial locations, visual features or objects. Imaging data provide support for all three types of visual selection, with attention modulating activity in areas specialized for processing the attended attributes.
How does attention affect the neural response to a stimulus? Attentional modulation of neural activity can reflect a multiplicative gain of the sensory response, or an additive increase in baseline activity. Although most imaging results are consistent with a gain mechanism, there is now evidence for top-down baseline shifts: attention can increase neural activity in extrastriate and even striate cortex in the absence of a stimulus.
Where do attentional signals come from? Recent evidence indicates that the source of attentional modulation stems from a fronto-parietal attention network. Several areas of this network participate in many kinds of attentional processing, including spatial orienting, eye movements, nonspatial selection and attention in non-visual modalities.
We are not passive recipients of the information that impinges on our retinae, but active participants in our own perceptual processes. Visual experience depends critically on attention. We select particular aspects of a visual scene for detailed analysis and control of subsequent behaviour, but ignore other aspects so completely that moments after they disappear from view we cannot report anything about them. Here we show that functional neuroimaging is revealing much more than where attention happens in the brain; it is beginning to answer some of the oldest and deepest questions about what visual attention is and how it works.
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We thank M. Chun, P. Downing, R. Epstein, Y. Jiang, M. Shuman and D. Somers for helpful comments on the manuscript. Work on this paper was supported by a Human Frontiers grant to N.K.
- INATTENTIONAL BLINDNESS
In a typical experiment, the subject decides which is longer, the horizontal or vertical arm of a large cross presented at fixation. An unexpected stimulus is then presented in the region of the cross, and immediately after the subject responds to the cross they are asked if they saw anything else. On a substantial number of trials, subjects do not report noticing the presence of the object at all.
- EVENT-RELATED POTENTIALS
Electrical potentials generated in the brain as a consequence of synchronized activation of neuronal networks by external stimuli. These evoked potentials are recorded at the scalp and consist of precisely timed sequences of waves or `components' (Fig. 1).
- EXTRASTRIATE CORTEX
All visually responsive areas of cortex except primary visual cortex.
- PRIMARY VISUAL CORTEX
The cortical area that is the main recipient of visual information coming from the retinae (by way of the lateral geniculate nucleus, or LGN); also known as V1 or striate cortex.
- VENTRAL AND DORSAL VISUAL PATHWAYS
Visual information coming from V1 is processed in two interconnected but partly dissociable visual pathways, a `ventral' pathway extending into the temporal lobe thought to be primarily involved in visual object recognition, and a `dorsal' pathway extending into the parietal lobes thought to be more involved in extracting information about `where' an object is or `how' to execute visually guided action towards it.
- RETINOTOPIC MAPPING
An fMRI procedure in which the borders of retinotopic visual areas (V1, V2, V3, and so on) are delineated, along with a representation of eccentricity and polar angle.
- BOTTOM-UP, FEEDFORWARD PROCESSING
Information processing that proceeds in a single direction from sensory input, through perceptual analysis, towards motor output, without involving feedback information flowing backwards from `higher' centres to `lower' centres.
- TOP-DOWN FEEDBACK
The flow of information from `higher' to `lower' centres.
Middle temporal and medial superior temporal extrastriate areas involved in the analysis of visual motion information.
- FUSIFORM FACE AREA (FFA)
A cortical region in the middle fusiform gyrus that responds at least twice as strongly in fMRI when subjects view faces as when they view various nonface stimuli.
- PARAHIPPOCAMPAL PLACE AREA
A bilateral region in parahippocampal cortex that produces at least twice as strong a signal in fMRI when subjects view images of places (including indoor and outdoor scenes and houses) as when they view images of nonplaces (for example, objects and faces).
- BASELINE SHIFTS
The increased response in a given neural population in an attended compared with unattended condition when no stimulus is present at all. Such effects imply that attention increases neural activity in an additive rather than a multiplicative fashion. That is, the magnitude of the response to a given stimulus when attended (A) should be higher by a constant K than the magnitude of the response to the same stimulus when unattended ( U), or U + K = A.
- GAIN MODULATION
The multiplicatively higher response to an attended compared with an unattended stimulus. If attention works by gain modulation then Ug = A, that is, the magnitude of the response to a given stimulus when attended ( A) should equal the product of an attentional gain multiplier (g) and the magnitude of response to the same stimulus when unattended (U).
A neurological syndrome (often involving damage to right parietal cortex) in which patients show a marked difficulty in the ability to detect or respond to information in the contralesional field.
In displays composed of identical distractor stimuli (for example, red Xs), a stimulus with a unique feature (for example, a blue X) can be detected rapidly and effortlessly, with little or no increase in reaction time as the number of distractor stimuli increases.
Lateral intraparietal area in the posterior parietal cortex of the monkey; single-unit physiological studies have shown that this area contains visually sensitive cells that increase their firing rate when a stimulus in their receptive field is attended, or is a target for a stimulus-driven or memory-guided saccade.
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Kanwisher, N., Wojciulik, E. Visual attention: Insights from brain imaging. Nat Rev Neurosci 1, 91–100 (2000). https://doi.org/10.1038/35039043
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