Working memory (WM) involves maintaining information in an on-line state. One emerging view is that information in WM is maintained via sensory recruitment, such that information is stored via sustained activity in the sensory areas that encode the to-be-remembered information. Using functional magnetic resonance imaging, we observed that key sensory regions such as primary visual cortex (V1) showed little evidence of sustained increases in mean activation during a WM delay period, though such amplitude increases have typically been used to determine whether a region is involved in on-line maintenance. However, a multivoxel pattern analysis of delay-period activity revealed a sustained pattern of activation in V1 that represented only the intentionally stored feature of a multifeature object. Moreover, the pattern of delay activity was qualitatively similar to that observed during the discrimination of sensory stimuli, suggesting that WM representations in V1 are reasonable "copies" of those evoked during pure sensory processing.Working memory (WM) allows the on-line storage of behaviorally relevant information. One emerging view is that WM is supported by the same neural mechanisms that encode the sensory information being remembered (we term this the sensory-recruitment model of WM; see Awh & Jonides, 2001;D'Esposito, 2007;Jonides, Lacey, & Nee, 2005;Postle, 2006). For example, neurons in face-selective regions of inferotemporal cortex show sustained amplitude increases while an observer is holding a face in WM (Chelazzi, Miller, Duncan, & Desimone, 1993;Courtney, Ungerleider, Keil, & Haxby, 1997;Druzgal & D'Esposito, 2001;Lepsien & Nobre, 2007;Miller, Li, & Desimone, 1993;Ranganath, Cohen, Dam, & D'Esposito, 2004). The sensory-recruitment hypothesis assumes that this activity represents the specific stimulus values of the stored items. Here we report a study in which functional magnetic resonance imaging (fMRI) and multivoxel pattern analysis (MVPA) provided direct support for this claim, showing that activation patterns in relevant sensory regions represent the specific stimulus value that is held in WM.MVPA provides a useful tool for identifying the neural regions that mediate WM by focusing on changes in activation patterns as opposed to simply changes in the mean amplitude of the blood-oxygenation-level-dependent (BOLD) response. For example, Offen, Schluppeck, and Heeger (in press) used fMRI to index activation changes in primary visual cortex (V1), a region known to represent orientation and spatial frequency. Although mean response amplitudes in V1 increased during sustained deployments of spatial attention, activation levels were indistinguishable from a low-level baseline when information about orientation (or spatial frequency) was stored in WM. This finding appears to contradict the sensory-recruitment model. preferentially to the remembered orientation should become more active, whereas neurons tuned away from the remembered orientation should be suppressed (relatively speaking; see, e.g.,...
Summary Working memory (WM) enables the storage and manipulation of information in an active state. WM storage has long been associated with sustained increases in activation across a network of frontal and parietal cortical regions. However, recent evidence suggests that these regions primarily encode information related to general task goals rather than feature-selective representations of specific memoranda. These goal-related representations are thought to provide top-down feedback that coordinates the representation of fine-grained details in early sensory areas. Here, we test this model using fMRI-based reconstructions of remembered visual details from region-level activation patterns. We could reconstruct high-fidelity representations of a remembered orientation based on activation patterns in occipital visual cortex and in several sub-regions of frontal and parietal cortex, independent of sustained increases in mean activation. These results challenge models of WM that postulate disjoint frontoparietal “top-down control” and posterior sensory “feature storage” networks.
Working memory (WM) enables the storage and manipulation of limited amounts of information over short periods. Prominent models posit that increasing the number of remembered items decreases the spiking activity dedicated to each item via mutual inhibition, which irreparably degrades the fidelity of each item’s representation. We tested these models by determining if degraded memory representations could be recovered following a post-cue indicating which of several items in spatial WM would be recalled. Using an fMRI-based image reconstruction technique, we identified impaired behavioral performance and degraded mnemonic representations with elevated memory load. However, in several cortical regions, degraded mnemonic representations recovered substantially following a post-cue, and this recovery tracked behavioral performance. These results challenge pure spike-based models of WM and suggest that remembered items are additionally encoded within latent or hidden neural codes that can help reinvigorate active WM representations.
Summary Working memory (WM) enables the maintenance and manipulation of information relevant to behavioral goals. Variability in WM ability is strongly correlated with IQ [1] and WM function is impaired in many neurological and psychiatric disorders [2, 3], suggesting that this system is a core component of higher cognition. WM storage is thought to be mediated by patterns of activity in neural populations selective for specific properties (e.g., color, orientation, location, motion direction) of memoranda [4–13]. Accordingly, many models propose that differences in the amplitude of these population responses should be related to differences in memory performance [14, 15]. Here, we used functional magnetic resonance imaging (fMRI) and an image reconstruction technique based on a spatial encoding model [16] to visualize and quantify population-level memory representations supported by multivoxel patterns of activation within regions of occipital, parietal and frontal cortex while participants precisely remembered the location(s) of 0, 1, or 2 small stimuli. We successfully reconstructed images containing representations of the remembered – but not forgotten – locations within regions of occipital, parietal, and frontal cortex using delay-period activation patterns. Critically, the amplitude of representations of remembered locations and behavioral performance both decreased with increasing memory load. These results suggest that differences in visual WM performance between memory load conditions are mediated by changes in the fidelity of large-scale population response profiles distributed across multiple areas of human cortex.
Recent studies suggest that the temporary storage of visual detail in working memory is mediated by sensory recruitment or sustained patterns of stimulus-specific activation within feature-selective regions of visual cortex. According to a strong version of this hypothesis, the relative “quality” of these patterns should determine the clarity of an individual’s memory. Here, we provide a direct test of this claim. We used fMRI and a forward encoding model to characterize population-level orientation-selective responses in visual cortex while human participants held an oriented grating in memory. This analysis, which enables a precise quantitative description of multivoxel, population-level activity measured during working memory storage, revealed graded response profiles whose amplitudes were greatest for the remembered orientation and fell monotonically as the angular distance from this orientation increased. Moreover, interparticipant differences in the dispersion—but not the amplitude—of these response profiles were strongly correlated with performance on a concurrent memory recall task. These findings provide important new evidence linking the precision of sustained population-level responses in visual cortex and memory acuity.
Recent studies suggest that visual features are stored in working memory (WM) via sensory recruitment or sustained stimulus-specific patterns of activity in cortical regions that encode memoranda. One important question concerns the spatial extent of sensory recruitment. One possibility is that sensory recruitment is restricted to neurons that are retinotopically mapped to the positions occupied by the remembered items. Alternatively, specific feature values could be represented via a spatially global recruitment of neurons that encode the remembered feature, regardless of the retinotopic position of the remembered stimulus. Here, we evaluated these alternatives by requiring subjects to remember the orientation of a grating presented in the left or right visual field. Functional magnetic resonance imaging and multivoxel pattern analysis were then used to examine feature-specific activations in early visual regions during memory maintenance. Activation patterns that discriminated the remembered feature were found in regions of contralateral visual cortex that corresponded to the retinotopic position of the remembered item, as well as in ipsilateral regions that were not retinotopically mapped to the position of the stored stimulus. These results suggest that visual details are held in WM through a spatially global recruitment of early sensory cortex. This spatially global recruitment may enhance memory precision by facilitating robust population coding of the stored information.
Spatial attention has been postulated to facilitate perceptual processing via several different mechanisms. For instance, attention can amplify neural responses in sensory areas (sensory gain), mediate neural variability (noise modulation), or alter the manner in which sensory signals are selectively read out by postsensory decision mechanisms (efficient readout). Even in the context of simple behavioral tasks, it is unclear how well each of these mechanisms can account for the relationship between attention-modulated changes in behavior and neural activity because few studies have systematically mapped changes between stimulus intensity, attentional focus, neural activity, and behavioral performance. Here, we used a combination of psychophysics, event-related potentials (ERPs), and quantitative modeling to explicitly link attention-related changes in perceptual sensitivity with changes in the ERP amplitudes recorded from human observers. Spatial attention led to a multiplicative increase in the amplitude of an early sensory ERP component (the P1, peaking ϳ80 -130 ms poststimulus) and in the amplitude of the late positive deflection component (peaking ϳ230 -330 ms poststimulus). A simple model based on signal detection theory demonstrates that these multiplicative gain changes were sufficient to account for attention-related improvements in perceptual sensitivity, without a need to invoke noise modulation. Moreover, combining the observed multiplicative gain with a postsensory readout mechanism resulted in a significantly poorer description of the observed behavioral data. We conclude that, at least in the context of relatively simple visual discrimination tasks, spatial attention modulates perceptual sensitivity primarily by modulating the gain of neural responses during early sensory processing
Are resources in visual working memory allocated in a continuous or a discrete fashion? On the one hand, flexible resource models suggest that capacity is determined by a central resource pool that can be flexibly divided such that items of greater complexity receive a larger share of resources. On the other hand, if capacity in working memory is defined in terms of discrete storage "slots" then observers may be able to determine which items are assigned to a slot but not how resources are divided between stored items. To test these predictions, we manipulated the relative complexity of the items to be stored while holding the number items constant. Although mnemonic resolution declined when set size increased (Experiment 1), resolution for a given item was unaffected by large variations in the complexity of the other items to be stored when set size was held constant (Experiments 2-4). Thus, resources in visual working memory are distributed in a discrete slot-based fashion, even when inter-item variations in complexity motivate an asymmetrical division of resources across items.
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