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Review
. 2021 Sep 15:7:257-277.
doi: 10.1146/annurev-vision-032321-100012. Epub 2021 Jul 9.

Visual Remapping

Affiliations
Review

Visual Remapping

Julie D Golomb et al. Annu Rev Vis Sci. .

Abstract

Our visual system is fundamentally retinotopic. When viewing a stable scene, each eye movement shifts object features and locations on the retina. Thus, sensory representations must be updated, or remapped, across saccades to align presaccadic and postsaccadic inputs. The earliest remapping studies focused on anticipatory, presaccadic shifts of neuronal spatial receptive fields. Over time, it has become clear that there are multiple forms of remapping and that different forms of remapping may be mediated by different neural mechanisms. This review attempts to organize the various forms of remapping into a functional taxonomy based on experimental data and ongoing debates about forward versus convergent remapping, presaccadic versus postsaccadic remapping, and spatial versus attentional remapping. We integrate findings from primate neurophysiological, human neuroimaging and behavioral, and computational modeling studies. We conclude by discussing persistent open questions related to remapping, with specific attention to binding of spatial and featural information during remapping and speculations about remapping's functional significance.

Keywords: attention; eye movements; retinotopic; saccade; spatiotopic; visual perception.

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Figures

Figure 1:
Figure 1:. Saccades can disrupt both visual and attentional representations in retinotopic brain areas.
Pre-saccadic (A) and post-saccadic (B) representations of a complex visual scene by a population of retinotopic visual neurons. Black plus indicates the current fixation position, arrow indicates the upcoming saccade, and the colored dashed circles indicate the neurons’ spatial receptive fields. Before the saccade, the purple neuron represents the small distant child, but after the saccade the same object is now represented by the red neuron, while the purple neuron represents a different object. In the context of attention, if spatial attention is directed to the small child before the saccade, facilitating activity in the purple neuron, then in the absence of some form of remapping, attention will be mislocalized to the wrong target (trees) after the saccade.
Figure 2:
Figure 2:. Forward vs convergent remapping.
(A-B) Single neuron schematic for forward spatial remapping (A) and convergent remapping (B). Black dot is initial fixation target (current gaze position), and green dot is saccade target. Black dashed circle depicts the neuron’s current receptive field (RF), red dashed circle indicates the forward remapping field location (future field), and purple indicates the convergent remapping field location (toward saccade target). (C) Time course of forward spatial remapping from a retinal perspective, with schematic centered on current gaze position at each time point. (Note Panel A depicts remapping instead from a world-centered perspective.) Timeframes illustrate how the retinal RF position changes under the influence of the saccade plan, shifting to a new retinal location during pre-saccadic remapping, and returning to the classical RF position once fixation is acquired at the saccade target (green). Yellow shading indicates the fovea and parafoveal area of the retina. (D) Convergent spatial remapping can enhance processing at saccade endpoints. All RFs shift towards the saccade target during remapping, resulting in an increase in the density of the neural representation around the saccade target that can mimic attentional gain effects.
Figure 3.
Figure 3.. Predictive Remapping vs Retinotopic Attentional Trace.
(A) Hypothetical responses of two visual neurons with different spatial receptive fields. The beige interval indicates the period prior to saccade initiation, the blue interval after. Yellow circle represents to-be-attended spatiotopic location. Before the saccade, the attended location falls within Neuron A’s receptive field; after the saccade, it falls in Neuron B’s. ‘Predictive remapping’ is when Neuron B begins to respond in anticipation of the saccade. ‘Retinotopic attentional trace’ is when Neuron A continues to respond for a period of time after the eye movement. Thus, there is a period of time where both spatiotopic and retinotopic locations are facilitated. (B) Corresponding locations for a behavioral study. Figure adapted from Golomb 2019. (Note to Annual Reviews: I am an author of this article, and the publisher has confirmed that I retain the rights to re-publish it.)
Figure 4.
Figure 4.. Spatial receptive field remapping vs attentional pointer remapping.
Left side: Schematic illustrating remapping mechanism where spatial receptive fields (RFs) transiently shift in anticipation of the saccade. Right side: Schematic illustrating alternative attentional pointer remapping mechanism. Gray boxes illustrate the locations of 3 neurons’ RFs (dashed circles), with current gaze position at the black dot, and the green dot representing the saccade target. The orange star indicates the stimulus probe, and the yellow shading indicates the attentional focus. Next to each box is a simplified diagram of the same 3 neurons (conceptualized here as LIP neurons), and the corresponding V1 neurons feeding feed-forward input (blue arrows indicate these connections). During the initial fixation period (top row), the stimulus falls in the RF of neuron 2. After the saccade (bottom row), the stimulus falls in the RF of neuron 1. During remapping (middle row), neuron 1 becomes active in anticipation of the saccade. The spatial RF remapping mechanism says this is because the RFs shift spatially to their future fields, which could be conceptualized as a remapping of which retinotopic V1 neurons feed into the LIP neurons, such that the neurons become transiently sensitive to a different portion of the visual field. The attentional pointer mechanism instead says that the RFs remain veridical, but the new set of neurons becomes facilitated in anticipation (i.e., the attention pointer remaps from neuron 2 to neuron 1). In both cases, the remapping signal could come from corollary discharge signals from an area such as thalamic MD.

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