Dr. rer. nat. Thomas Christophel
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Forschungsthemen1
Adaptive kortikal Organisation im Dienste der verteilten Arbeitsgedächtnisspeicherung
Quelle ↗Förderer: DFG Nachwuchsgruppe Zeitraum: 01/2021 - 12/2026 Projektleitung: Dr. rer. nat. Thomas Christophel
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Publikationen25
Top 25 nach Zitationen — Quelle: OpenAlex (BAAI/bge-m3 embedded für Matching).
Trends in Cognitive Sciences · 961 Zitationen · DOI
Journal of Neuroscience · 323 Zitationen · DOI
How content is stored in the human brain during visual short-term memory (VSTM) is still an open question. Different theories postulate storage of remembered stimuli in prefrontal, parietal, or visual areas. Aiming at a distinction between these theories, we investigated the content-specificity of BOLD signals from various brain regions during a VSTM task using multivariate pattern classification. To participate in memory maintenance, candidate regions would need to have information about the different contents held in memory. We identified two brain regions where local patterns of fMRI signals represented the remembered content. Apart from the previously established storage in visual areas, we also discovered an area in the posterior parietal cortex where activity patterns allowed us to decode the specific stimuli held in memory. Our results demonstrate that storage in VSTM extends beyond visual areas, but no frontal regions were found. Thus, while frontal and parietal areas typically coactivate during VSTM, maintenance of content in the frontoparietal network might be limited to parietal cortex.
Nature Neuroscience · 234 Zitationen · DOI
NeuroImage · 105 Zitationen · DOI
Visual Cognition · 80 Zitationen · DOI
Visual working memory (VWM) relies on a distributed cortical network. Yet, the extent to which individual cortical areas, like early visual cortex and intraparietal sulcus, are essential to VWM storage remains debated. Here, we reanalyze key datasets from two independent labs to address three topics at the forefront of current-day VWM research: Resiliency of mnemonic representations against visual distraction, the role of attentional priority in memory, and brain–behavior relationships. By utilizing different analysis approaches, each designed to test different aspects of mnemonic coding, our results provide a comprehensive perspective on the role of early visual and intraparietal areas. We emphasize the importance of analysis choices, and how a thorough understanding of the principles they test is crucial for unraveling the distributed mechanisms of VWM. Consequently, we caution against the idea of a singular essential storage area, which could limit our comprehension of the VWM system.
Journal of Neuroscience · 79 Zitationen · DOI
Visual working memory (VWM) is used to maintain visual information available for subsequent goal-directed behavior. The content of VWM has been shown to affect the behavioral response to concurrent visual input, suggesting that visual representations originating from VWM and from sensory input draw upon a shared neural substrate (i.e., a <i>sensory recruitment</i> stance on VWM storage). Here, we hypothesized that visual information maintained in VWM would enhance the neural response to concurrent visual input that matches the content of VWM. To test this hypothesis, we measured fMRI BOLD responses to task-irrelevant stimuli acquired from 15 human participants (three males) performing a concurrent delayed match-to-sample task. In this task, observers were sequentially presented with two shape stimuli and a retro-cue indicating which of the two shapes should be memorized for subsequent recognition. During the retention interval, a task-irrelevant shape (the probe) was briefly presented in the peripheral visual field, which could either match or mismatch the shape category of the memorized stimulus. We show that this probe stimulus elicited a stronger BOLD response, and allowed for increased shape-classification performance, when it matched rather than mismatched the concurrently memorized content, despite identical visual stimulation. Our results demonstrate that VWM enhances the neural response to concurrent visual input in a content-specific way. This finding is consistent with the view that neural populations involved in sensory processing are recruited for VWM storage, and it provides a common explanation for a plethora of behavioral studies in which VWM-matching visual input elicits a stronger behavioral and perceptual response.<b>SIGNIFICANCE STATEMENT</b> Humans heavily rely on visual information to interact with their environment and frequently must memorize such information for later use. Visual working memory allows for maintaining such visual information in the mind's eye after termination of its retinal input. It is hypothesized that information maintained in visual working memory relies on the same neural populations that process visual input. Accordingly, the content of visual working memory is known to affect our conscious perception of concurrent visual input. Here, we demonstrate for the first time that visual input elicits an enhanced neural response when it matches the content of visual working memory, both in terms of signal strength and information content.
NeuroImage · 70 Zitationen · DOI
Journal of Magnetic Resonance Imaging · 35 Zitationen · DOI
1 Technical Efficacy: Stage 1 J. MAGN. RESON. IMAGING 2017;46:134-141.
Human Brain Mapping · 31 Zitationen · DOI
It has been suggested that visual images are memorized across brief periods of time by vividly imagining them as if they were still there. In line with this, the contents of both working memory and visual imagery are known to be encoded already in early visual cortex. If these signals in early visual areas were indeed to reflect a combined imagery and memory code, one would predict them to be weaker for individuals with reduced visual imagery vividness. Here, we systematically investigated this question in two groups of participants. Strong and weak imagers were asked to remember images across brief delay periods. We were able to reliably reconstruct the memorized stimuli from early visual cortex during the delay. Importantly, in contrast to the prediction, the quality of reconstruction was equally accurate for both strong and weak imagers. The decodable information also closely reflected behavioral precision in both groups, suggesting it could contribute to behavioral performance, even in the extreme case of completely aphantasic individuals. Our data thus suggest that working memory signals in early visual cortex can be present even in the (near) absence of phenomenal imagery.
Cerebral Cortex · 30 Zitationen · DOI
Traditional views of visual working memory postulate that memorized contents are stored in dorsolateral prefrontal cortex using an adaptive and flexible code. In contrast, recent studies proposed that contents are maintained by posterior brain areas using codes akin to perceptual representations. An important question is whether this reflects a difference in the level of abstraction between posterior and prefrontal representations. Here, we investigated whether neural representations of visual working memory contents are view-independent, as indicated by rotation-invariance. Using functional magnetic resonance imaging and multivariate pattern analyses, we show that when subjects memorize complex shapes, both posterior and frontal brain regions maintain the memorized contents using a rotation-invariant code. Importantly, we found the representations in frontal cortex to be localized to the frontal eye fields rather than dorsolateral prefrontal cortices. Thus, our results give evidence for the view-independent storage of complex shapes in distributed representations across posterior and frontal brain regions.
NeuroImage · 22 Zitationen · DOI
Working memory contents are represented in neural activity patterns across multiple regions of the cortical hierarchy. A division of labor has been proposed where more anterior regions harbor increasingly abstract and categorical representations while the most detailed representations are held in primary sensory cortices. Here, using fMRI and multivariate encoding modeling, we demonstrate that for color stimuli categorical codes are already present at the level of extrastriate visual cortex (V4 and VO1), even when subjects are neither implicitly nor explicitly encouraged to categorize the stimuli. Importantly, this categorical coding was observed during working memory, but not during perception. Thus, visual working memory is likely to rely at least in part on categorical representations. SIGNIFICANCE STATEMENT: Working memory is the representational basis for human cognition. Recent work has demonstrated that numerous regions across the human brain can represent the contents of working memory. We use fMRI brain scanning and machine learning methods to demonstrate that different regions can represent the same content differently during working memory. Reading out the neural codes used to store working memory contents, we show that already in sensory cortex, areas V4 and VO1 represent color in a categorical format rather than a purely sensory fashion. Thereby, we provide a better understanding of how different regions of the brain might serve working memory and cognition.
Psychological Science · 21 Zitationen · DOI
When looking at static visual images, people often exhibit mental animation, anticipating visual events that have not yet happened. But what determines when mental animation occurs? Measuring mental animation using localized brain function (visual motion processing in the middle temporal and middle superior temporal areas, MT+), we demonstrated that animating static pictures of objects is dependent both on the functionally relevant spatial arrangement that objects have with one another (e.g., a bottle above a glass vs. a glass above a bottle) and on the linguistic judgment to be made about those objects (e.g., "Is the bottle above the glass?" vs. "Is the bottle bigger than the glass?"). Furthermore, we showed that mental animation is driven by functional relations and language separately in the right hemisphere of the brain but conjointly in the left hemisphere. Mental animation is not a unitary construct; the predictions humans make about the visual world are driven flexibly, with hemispheric asymmetry in the routes to MT+ activation.
Cortex · 17 Zitationen · DOI
NeuroImage · 13 Zitationen · DOI
Representations of sensory working memory can be found across the entire neocortex. But how are verbal working memory (VWM) contents retained in the human brain? Here we used fMRI and multi-voxel pattern analyses to study Chinese native speakers (15 males, 13 females) memorizing Chinese characters. Chinese characters are uniquely suitable to study VWM because verbal encoding is encouraged by their complex visual appearance and monosyllabic pronunciation. We found that activity patterns in Broca's area and left premotor cortex carried information about the memorized characters. These language-related areas carried (1) significantly more information about cued characters than those not cued for memorization, (2) significantly more information on the left than the right hemisphere and (3) significantly more information about Chinese symbols than complex visual patterns which are hard to verbalize. In contrast, early visual cortex carries a comparable amount of information about cued and uncued stimuli and is thus unlikely to be involved in memory retention. This study provides evidence for verbal working memory maintenance in a distributed network of language-related brain regions, consistent with distributed accounts of WM. The results also suggest that Broca's area and left premotor cortex form the articulatory network which serves articulatory rehearsal in the retention of verbal working memory contents.
NeuroImage · 11 Zitationen · DOI
Visual working memory (VWM) allows for keeping visual information available for upcoming goal-directed behavior, while new visual input is processed concurrently. Interactions between the mnemonic and perceptual systems cause VWM to affect the processing of visual input in a content-specific manner: visual input that is initially suppressed from consciousness is detected faster when it matches rather than mismatches the content of VWM. It is currently under debate whether such mnemonic influences on perception occur prior to or after conscious access. To address this issue, we investigated whether VWM content modulates the neural response to visual input that remains suppressed from consciousness. We measured fMRI responses to interocularly suppressed stimuli in 20 human participants performing a delayed match-to-sample task: Participants were retro-cued to memorize one of two geometrical shapes for subsequent recognition. During retention, an interocularly suppressed peripheral stimulus (the probe) was briefly presented, which was either of the cued (memorized) or uncued (not memorized) shape category. We found no evidence that VWM content modulated the neural response to the probe. Substantial evidence for the absence of this modulation was found despite leveraging a highly liberal analysis approach: (1) selecting regions of interest that were particularly prone to detecting said modulation, and (2) using directional Bayesian tests favoring the presence of the hypothesized modulation. We did observe faster detection of memory-matching compared to memory-mismatching probes in a behavioral control experiment, thus validating the stimulus set. We conclude that VWM impacts the processing of visual input only once suppression is mostly alleviated.
bioRxiv (Cold Spring Harbor Laboratory) · 8 Zitationen · DOI
Abstract It has been suggested that visual images are memorized across brief periods of time by vividly imagining them as if they still were there. In line with this, the contents of both working memory and visual imagery are known to be encoded already in early visual cortex. If these signals in early visual areas were indeed to reflect a combined imagery and memory code, one would predict them to be weaker for individuals with reduced visual imagery vividness. Here, we systematically investigated this question in two groups of participants. Strong and weak imagers were asked to remember images across brief delay periods. We were able to reliably reconstruct the memorized stimuli from early visual cortex during the delay. Importantly, in contrast to the prediction, the quality of reconstruction was equally accurate for both strong and weak imagers. The decodable information also closely reflected behavioral precision in both groups, suggesting it could contribute to behavioral performance, even in the extreme case of completely aphantasic individuals. Our data thus suggest that working memory signals in early visual cortex can be present even in the (near) absence of phenomenal imagery.
Journal of Cognition · 7 Zitationen · DOI
Visual working memory and verbal storage are often investigated independently of one another. However, a growing body of evidence suggests that naming visual stimuli can provide an advantage in performance during visual working memory tasks. On the other hand, there is also evidence that labeling could lead to biases in recall. Here, we present an exploratory investigation of verbal labels associated with the memorization of simple visuo-spatial stimuli, and how the use of these labels informs recall behavior of the same stimuli in a separate working memory task. English-speaking participants performed a working memory task with orientation and location stimuli, followed by a separate naming task featuring the same stimuli. We found a diverse set of labels employed frequently and with a consistent distribution across stimulus types, the stimulus space, and among participants. The use of individual spatial words, predicted class 1 cardinal biases in memory (i.e. the observation that cardinal stimuli are more accurately recalled than non-cardinal ones). Conversely, words expressing uncertainty (e.g. 'slightly', 'near') predicted class 2 cardinal bias (i.e. recall biases away from the cardinal planes). This relationship between word use and recall biases is consistent with shared representational resources that are used for both visuo-spatial and verbal working memory.
bioRxiv (Cold Spring Harbor Laboratory) · 5 Zitationen · DOI
Abstract While distributed cortical areas represent working memory contents, their necessity for memory maintenance has been questioned. We examined the differential effects of maintaining multiple items on the neural information across cortical regions. In each trial of the fMRI experiment, participants (n=81) had to memorize two items, each either an orientation or a pure pitch, for 13.8s and continuously recalled the target after the delay. We kept the overall working memory load constant, but varied the sensory modality of each item to vary the effective visual load. We show that increasing visual load decreased behavioural recall performance for orientations and continuous orientation-specific decodable information in visual cortex but less so infrontoparietal areas. Simulations show that this drop in decodable information is best interpreted as a drop in mnemonic information represented by patterns of visual cortex activity. Our results provide evidence for shared labour of visual cortices, where maintaining two versus one orientation leads to a loss in representational fidelity, and anterior cortices, where multiple items could be represented in a more robust but less precise format.
Journal of Vision · 4 Zitationen · DOI
Distributed Visual Working Memory Stores Revealed by Multivariate Pattern Analyses Thomas B. Christophel, Chang Yan, Carsten Allefeld & John-Dylan Haynes The storage buffers retaining visual working memory contents were originally postulated to reside in prefrontal cortex. Recently, a dissenting view has evolved claiming that working memory content depends on distributed storage in sensory brain regions. We provide strong evidence for this claim in a series of fMRI experiments investigating the content-specificity of delay-period activity using multivariate pattern analyses. Representations of color and motion patterns as well as complex shapes were identified in early visual, and lateral occipital posterior parietal cortex, but also in the frontal eye fields. A meta-analysis of content-specificity within these brain areas revealed large inter-areal differences critically depending on whether the stimuli were smooth global patterns or shapes with clear edges and on whether stimuli varied across color, luminance or motion direction dimensions. In addition, we show that areas beyond early visual cortex retain information in an inherently view-independent format and that coding of a given stimulus in higher visual areas is not solely driven by the visual display originally shown. Instead, the representation changes when a subject mentally transforms what they are holding in mind (i.e. during mental rotation). Extending our findings on visual working memory, we show that verbal content (Chinese Characters memorized by native speakers of Chinese) is selectively stored in prefrontal areas, more specifically Broca’s area and articulatory premotor cortex. Finally, while working memory storage seems to be represented in a distributed way, working memory control could be traced to dorsolateral prefrontal cortex regardless of what content was memorized.
Journal of Cognitive Neuroscience · 2 Zitationen · DOI
Items held in visual working memory can be quickly updated, replaced, removed, and even manipulated in accordance with current behavioral goals. Here, we use multivariate pattern analyses to identify the patterns of neuronal activity that realize the executive control processes supervising these flexible stores. We find that portions of the middle temporal gyrus and the intraparietal sulcus represent what item is cued for continued memorization independently of representations of the item itself. Importantly, this selection-specific activity could not be explained by sensory representations of the cue and is only present when control is exerted. Our results suggest that the selection of memorized items might be controlled in a distributed and decentralized fashion. This evidence provides an alternative perspective to the notion of "domain general" central executive control over memory function.
Journal of Vision · 2 Zitationen · DOI
Visual working memory (VWM) relies on a distributed cortical network. Yet, the role of individual cortical areas, like early visual cortex (EVC) and intraparietal sulcus (IPS), remains debated. Criteria have been suggested to determine if an area is essential for storage, such as resiliency against visual distraction, and correlations with behavior. Here, we reanalyzed existing data from two independent labs, and caution that adherence to simple criteria could limit our comprehension of the VWM system. Instead, we encourage close consideration of analysis choices for a more complete perspective. When participants remembered an orientation while simultaneously viewing different visual distractors (Rademaker et al., 2019) fMRI activity patterns in EVC and IPS did not distinguish between distractor conditions. While such resiliency implies that both areas are critical for VWM storage, the analysis used (leave-one-out cross-validation) capitalizes on any signal differentiating memory representations during the delay. Instead, an analysis using sensory driven responses for model-training captures only representations that are “sensory-like”, and can yield different conclusions. That analysis choices matter is further illustrated by a task with two memory items – one attended, one unattended (Christophel et al., 2018). Originally, delay-period representations of unattended items were not found in EVC. Our reanalysis reveals that with a model trained on stronger signals (the attended instead of the unattended items) EVC does represent unattended memory items. Finally, both datasets reveal a brain-behavior relationship in EVC, but not IPS. Before declaring EVC essential to storage, a careful examination of analysis choices (like the quantification and read-out of neural error) should be performed to guide interpretation. In sum, a thorough understanding of analyses and the specific principles they test is crucial for unraveling the mechanisms of VWM.
Journal of Vision · 2 Zitationen · DOI
Visual shape recognition can exhibit considerable invariance across changes in visual appearance and viewing position (Cichy, Chen, & Haynes, 2011; Edelman, 1997; Grill-Spector, Kourtzi, & Kanwisher, 2001). This raises the question whether also the retention of shape information in visual working memory (Baddeley & Hitch, 1974) exhibits such invariance. Here, we specifically investigated whether objects memorized across brief delays are encoded using a rotation-invariant code. While positioned in an MRI scanner, 22 healthy subjects memorized simple shape stimuli. To prevent subjects from using a semantic code, we chose abstract decagons as stimuli. These are randomly generated ten-sided shapes. In each trial, subjects had to memorize one shape indicated by a retro-cue method controlling for perceptional confounds (see Harrison & Tong, 2009). After a delay of 10 seconds, subjects had to identify which of two test decagons had a more similar shape (see Christophel, Hebart, & Haynes, 2012). Importantly, sample and test decagons were shown in random 2D-rotations to encourage the use of rotation-invariant representations. We used fMRI in combination with time-resolved multivariate searchlight decoding to identify areas that maintained the memorized object during the delay period using a rotation-invariant code (Cichy et al., 2011; Haynes & Rees, 2006; Kriegeskorte, Goebel, & Bandettini, 2006). Testing for classifier generalization between different rotational views of the same shape, we identified three regions that showed significant (p(FWE) <0.05) memory-specific information bilaterally: Lateral occipito-temporal cortex, posterior parietal cortex and the human frontal eye fields (see Petit, Clark, Ingeholm, & Haxby, 1997). These results demonstrate that invariant shape-coding in working memory is prevalent in perception-driven areas across the brain (see Postle, 2006). Importantly, our findings demonstrate that invariant visual memory representations do not require higher-order dorso-lateral prefrontal areas. Meeting abstract presented at VSS 2013
bioRxiv (Cold Spring Harbor Laboratory) · 1 Zitationen · DOI
Abstract A single visual stimulus can elicit multiple concurrent representations throughout the cortex. We show that during a visual working memory task several cortical regions utilize categorical, semantic, and spatial representational formats to maintain visual stimuli in a robust fashion. We assessed the nature of orientation representations in an fMRI dataset of 40 participants performing an orientation memory task using multivariate encoding modelling. Our results show that orientation representations across the cortex form a gradient of abstraction, with more veridical representations in sensory areas and more abstract, categorical codes in anterior areas. We use cross-condition modelling to demonstrate shared neural codes between orientation and congruent verbal and location stimuli, used at different points during the trial. These findings provide evidence for a distributed account of working memory storage, where memory representations, distinct in nature and content, are present concurrently throughout the cortex.
Journal of Vision · 1 Zitationen · DOI
Research into the neural basis of working memory relies heavily on the use of multivariate decoding techniques to ascertain the presence of neural representations of working memory content. Recent work attempts to understand the tuning properties underlying cortical representations using encoding modelling. Encoding modelling aims to explain the multivariate patterns underlying mnemonic function using stimulus dependent regressors called basis functions. These basis functions are capable of explicitly modelling the similarity relationships between different stimuli analogous to tuning functions. Here, we aim to evaluate the use of encoding modelling for the quantification of mnemonic representations using simulated and real data. We make use of a recently developed flexible simulation toolbox to simulate patterned neural activity generated using different underlying voxel tuning distributions across a large number of possible experimental designs (designSim). We quantify the information content of these neural signals as the variance explained by a given encoding model using cvCrossMANOVA. We demonstrate that realistically modelled neural representation can be more reliably identified using encoding models with realistic similarity assumptions as compared to a simplistic classifier-like models. We show that estimates of representational similarity between two conditions (e.g., cross-classification accuracy, correlation, and variance explained between conditions) are strongly biased by the signal-to-noise ratio of individual representations and provide an SNR-independent measure of pattern similarity by comparing variance explained within and between conditions. Finally, we ask whether encoding modelling can ascertain the precise representational code used for memorization. We show that, consistent with prior work, stimulus-driven representation of memorized contents can be fitted and explained using a large variety of differently shaped encoding models such. This means that a reliable fit alone gives little insight into the representational geometry (‘feature fallacy’). Instead, we demonstrate that comparing the explained variance of two or more competing models allows to reliably identify the true model.
Journal of Vision · 1 Zitationen · DOI
Categorical biases influence the recall of visual stimuli during working memory. This effect is well documented for colors, which are easily categorized into named color categories (‘red’, ‘green’, ‘blue’), and behavioral evidence suggests that this categorization can bias delayed recall toward prototypical color hues. For orientations and locations, cardinal stimuli (i.e. ‘horizontal’ and ‘vertical’) are remembered with significantly higher accuracy than non-cardinal ones and recall of noncardinal stimuli is biased away from these cardinals. We have recently shown that this difference in recall between categories seems to be reflected by different labeling strategies for cardinal and noncardinal stimuli. Words describing cardinal stimuli were used more selectively, while terms for non-cardinal stimuli (i.e. ‘diagonal’) were used more liberally and broadly. This suggested that subjects used verbal or categorical encoding strategies which in turn resulted in biased recall. Here, we investigated whether these different behavioral encoding strategies for cardinal and non-cardinal stimuli are also evident in neural representations during working memory. Subjects performed a standard orientation and location delayed recall task in an MRI scanner with no additional instruction regarding the use of verbal or non-verbal strategies. After these tasks, we also recorded the brain activity evoked when reading and listening to a set of spatial words used frequently to describe orientation and location stimuli (e.g., ‘horizontal’, ‘vertical’). We find that encoding cardinal and noncardinal stimuli evokes subject-unique neural activity patterns in posterior parietal cortices which are replicated for orientation and location stimuli. These neural activity patterns can entail both univariate and multivariate changes in neural recruitment. These results suggest that cardinal biases for orientation and location stimuli are a result of shared categorical-spatial representations outside of visual cortex. This highlights how both sensory and more anterior regions in concert enable working memory behavior using sensory, categorical, and potentially even verbal codes.
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Identität, Organisation und Kontakt aus HU-FIS.
- Name
- Dr. rer. nat. Thomas Christophel
- Titel
- Dr. rer. nat.
- Fakultät
- Lebenswissenschaftliche Fakultät
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- Institut für Psychologie
- Arbeitsgruppe
- NWG Distributed Cognition and Memory
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