Biology subjects

Griffiths, B. J.

Publications and source records attributed to Griffiths, B. J..

15 recordsLinked to original sources

bioRxiv2026

Comparing Electrical and Ultrasound Transcutaneous Vagus Nerve Stimulation (taVNS) on Associative Memory

Associative memory, the ability to bind and retrieve relationships between unrelated elements, is a cornerstone of human cognition and a primary target for neurorehabilitation. Vagus nerve stimulation (VNS) has emerged as a promising method to modulate the locus coeruleus-norepinephrine (LC-NE) system and hippocampal-prefrontal circuits essential for memory. However, the comparative efficacy of non-invasive modalities such as electrical (E-taVNS) and the emerging field of ultrasound (U-taVNS) remains poorly understood in the context of active recall. In this study, participants performed a crossmodal video-word associative memory task before and after receiving either E-taVNS or U-taVNS in active and sham conditions. We investigated whether these modalities enhance cued recall accuracy and retrieval reaction time. Our results revealed that neither E-taVNS nor U-taVNS significantly improved recall accuracy. However, E-taVNS significantly accelerated response times specifically for correctly recalled items. These findings suggest that while taVNS may not increase the likelihood of recalling associative memories, electrical stimulation may enhances the efficiency in which we do so. These findings suggest that electrical taVNS is a viable tool for facilitating memory search processes, though further research is required to optimize ultrasound parameters and validate mechanistic pathways through physiological monitoring.

neuroscience↗

bioRxiv2026

Alpha oscillations track the projection of reactivated memories into conscious awareness

By definition, episodic memory is a conscious phenomenon. Memory traces reactivated by the hippocampus and reinstated in the sensory cortices need to enter conscious awareness for them to be re-experienced and overtly recalled. However, it remains unclear whether such reactivation in-and-of-itself ensures that memories will be overtly recalled. To investigate this, magnetoencephalography (MEG) recordings were analysed from thirty-one participants (18 female, 13 male) completing a video-word pair associates memory task. When combining linear classifiers and spectral analyses, sensory cortical reactivation could be observed without overt recall occurring, suggesting reactivation does not guarantee overt recall. Instead, overt recall was additively predicted by (i) an increase in reactivated representations rhythmically fluctuating within the alpha band, and (ii) a decrease in total sensory neocortical alpha power. These results are consistent with accounts which propose that reactivation benefits from desynchronising the network to provide representational space for stimulus-specific information, and/or amplifying stimulus-specific information above residual noise. Altogether, these results suggest that representational reactivation can occur without overt recall, and suggest a role for alpha oscillations in projecting internally-generated representations into conscious awareness.

neuroscience↗

bioRxiv2025

Rhythmic light stimulation elicits multiple concurrent neural responses that separably shape human perception

Rhythmic light stimulation offers solutions to innumerable cognitive and neurological disorders. However, like any neuromodulatory technique, responses to rhythmic light stimulation are highly variable, producing challenges in replicating lab-based studies and translating findings to the clinic. Across three MEG/EEG experiments, we show that this variability can, in part, be attributed to rhythmic light stimulation eliciting multiple, coexisting neural responses which have separable impacts on cognition. Specifically, we find that rhythmic light stimulation produces distinct neural responses at the fundamental (f) and second harmonic (2f) frequencies, and that these responses are differentially shaped by endogenous oscillatory dynamics that vary across participants. Importantly, these responses separably contribute to perception, with harmonic gamma-band responses supporting the representation of stimulus-specific information, and the phase of harmonic alpha-band responses causally contributing to near-threshold visual perception. We reproduce these effects across datasets, paradigms, and oscillatory bands, suggesting that the multiplex oscillatory responses elicited by rhythmic light stimulation are a robust and pervasive phenomenon. We propose that the complexity of neural responses to rhythmic stimulation can explain why there is substantial variability between studies using these techniques, and that understanding these complex responses may help advance neuromodulatory technologies for both fundamental and clinical neuroscience. Supplementary MaterialAll supplementary material can be found at the end of this document.

neuroscience↗

bioRxiv2025

Higher baseline alpha power is associated with faster responses in visual search

Visual search models have long emphasised that task-relevant items must be prioritized for optimal performance. While it is known that search efficiency also benefits from active distractor inhibition, the underlying neuronal mechanisms are debated. Neuronal alpha oscillations (7-14 Hz) have been associated with functional inhibition of cortical excitability, as well as distractor suppression in spatial attention and visual working memory tasks. We therefore hypothesised that alpha oscillations similarly support the deselection of distractors in visual search. Using Magnetoencephalography (MEG), we here show that high alpha power before the onset of a complex search display is associated with faster search performance. Crucially, we used a General Linear Model (GLM) approach to control for confounds between alpha power and task duration, ruling out that this result was merely driven by practice effects paired with increased fatigue over time. In addition to spontaneous oscillatory activity, we quantified the cortical excitability to colours of the search stimuli based on Rapid Invisible Frequency Tagging (RIFT) responses. In contrast to our initial hypothesis, increased pre-search alpha power did not correlate with the RIFT response, providing no direct evidence for feature-specific inhibition of distracting stimuli by alpha. Our findings challenge the traditional view of alpha oscillations reducing visual processing, showing instead that increased occipital alpha power can enhance performance in a visual task. We propose that the increase in alpha power may reflect increased top-down control supporting visual search

neuroscience↗

bioRxiv2024

Guided Visual Search is associated with a feature-based priority map in early visual cortex

Visual search models have long emphasised that task-relevant items must be prioritised for optimal performance. While it is known that search efficiency also benefits from active distractor inhibition, the underlying neuronal mechanisms are debated. Here, we used MEG in combination with Rapid Invisible Frequency Tagging (RIFT) to probe how neuronal excitability in early visual cortex is modulated during feature-guided visual search. Participants were instructed to indicate the presence or absence of a letter "T" presented amongst 16 and 32 "L"s. In the guided search condition, participants were informed about the colour of the "T" and could infer the colour of the irrelevant distractors. In the unguided search condition, the target colour was unknown. We found that guided search was associated with enhanced RIFT responses to the target colour, and decreased responses to the distractor colour compared to unguided search. These results conceptually replicated using both a conventional coherence approach, as well as with a General Linear Model approach based on a single-trial measure of the RIFT response. The present findings expand on previous reports based on electrophysiology and fMRI in humans and non-human primates by revealing that feature-guidance in visual search affects neuronal excitability as early as primary visual cortex.

neuroscience↗

bioRxiv2023

Imperceptible gamma-band sensory stimulation enhances episodic memory retrieval

Enhanced gamma activity (30-100Hz) coincides with the successful recall of episodic memories, but it remains unknown whether this oscillatory activity is a cause or a consequence of the retrieval process. To address this question, we asked human participants to complete a paired associates memory task while undergoing sensory stimulation (at 65Hz, 43.3Hz and 32.5Hz). We observed that 65Hz and 32.5Hz sensory stimulation enhances recall compared to a baseline condition without stimulation. No similar effect was observed following 43.3Hz stimulation. Notably, while almost all participants could perceive 32.5Hz and 43.3Hz sensory stimulation, only a small proportion of participants ([~]10%) could perceive the 65Hz visual flicker, suggesting 65Hz sensory stimulation acts as an imperceptible intervention to enhance recall. To understand the dual action of 65Hz and 32.5Hz sensory stimulation on recall, we built three pyramidal-interneuronal network gamma (PING) models and drove them using the same stimulation protocols as in the behavioural task. The behavioural results could be reproduced by stimulating an endogenous [~]32Hz oscillation, but not by stimulating an endogenous [~]65Hz oscillation nor by stimulating a network without an endogenous oscillation. These results suggest that imperceptible 65Hz sensory stimulation enhances recall by harmonically entraining an endogenous [~]32.5Hz oscillation. Based on these findings, we propose that "slow" gamma oscillations play a causal role in episodic memory retrieval.

neuroscience↗

bioRxiv2023

Memory reactivation of real-world spatial orientation revealed by human electrophysiology.

Memory consolidation relies on the reactivation of previous experiences during sleep. The precise interplay of sleep-related oscillations (slow oscillations, spindles and ripples) is thought to coordinate the information flow between relevant brain areas, with ripples mediating memory reactivation. However, in humans empirical evidence for a role of ripples in memory reactivation is lacking. Here, we investigated the relevance of sleep oscillations and specifically ripples for memory reactivation during human sleep using targeted memory reactivation (TMR). Intracranial electrophysiology in epilepsy patients and scalp EEG in healthy participants revealed that elevated levels of SO-spindle activity promoted the read-out of TMR induced memory reactivation. Importantly, spindle-locked ripples recorded intracranially from the medial temporal lobe were found to be instrumental for memory reactivation to unfold during non-rapid eye movement (NREM) sleep. Our findings establish ripples as key-oscillation in human systems consolidation and emphasize the importance of the coordinated interplay of the cardinal sleep oscillations.

neuroscience↗

bioRxiv2023

Electrophysiological signatures of veridical head direction in humans

Information about heading direction is critical for navigation as it provides the means to orient ourselves in space. However, given that veridical head direction signals require physical rotation of the head and most human neuroimaging experiments depend upon fixing the head in position, little is known about how the human brain is tuned to such heading signals. To address this, we asked fifty-two healthy participants undergoing simultaneous EEG and motion tracking recordings (split into two experiments) and ten patients undergoing simultaneous intracranial EEG and motion tracking recordings to complete a series of orientation tasks in which they made physical head rotations to target positions. We then used a series of forward encoding models and linear mixed-effects models to isolate electrophysiological activity that was specifically tuned to heading direction. We identified a robust posterior central signature that predicts changes in veridical head orientation after regressing out confounds including sensory input and muscular activity. Both source localisation and intracranial analysis implicated the medial temporal lobe as the origin of this effect. Subsequent analyses disentangled head direction signatures from signals relating to head rotation and those reflecting location-specific effects. Lastly, when directly comparing head direction and eye gaze-related tuning, we found that the brain maintains both codes while actively navigating, with stronger tuning to head direction in the medial temporal lobe. Together, these results reveal a taxonomy of population-level head direction signals within the human brain that is reminiscent of those reported in the single units of rodents.

neuroscience↗

bioRxiv2022

Reconstructing Spatio-Temporal Trajectories of Visual Object Memories in the Human Brain

Our understanding of how information unfolds when we recall events from memory remains limited. In this study, we investigate whether the reconstruction of visual object memories follows a backward trajectory along the ventral visual stream with respect to perception, such that their neural feature representations are gradually reinstated from late areas close to the hippocampus backwards to lower-level sensory areas. We use multivariate analyses of fMRI activation patterns to map high-and low-level features of the object memories onto the brain during retrieval, and EEG-fMRI fusion to track the temporal evolution of the reactivated patterns. Participants studied new associations between verbs and randomly paired object images in an encoding phase, and subsequently recalled the objects when presented with the corresponding verb cue. Decoding reactivated memory features from fMRI activity revealed that retrieval patterns were dominated by conceptual features, represented in comparatively late visual and parietal areas. Representational-similarity-based fusion then allowed us to map the EEG patterns that emerged at each given time point of a trial onto the spatially resolved fMRI patterns. This fusion suggests that memory reconstruction proceeds from anterior fronto-temporal to posterior occipital and parietal regions, partly in line with a semantic-to-perceptual gradient. A linear regression on the peak time points of reactivated brain regions statistically confirms that the temporal progression is reversed with respect to encoding in ventral stream areas. Together, the results shed light onto the spatio-temporal trajectories along which memories and their constituent features are reconstructed during associative retrieval. Significance statementIn our work, we combined EEG and fMRI to investigate which features of a visual object are reactivated when recalled from episodic memory, and how the memory reconstruction stream unfolds over time and across the brain. Our findings suggest that with respect to perception, memory retrieval follows a backwards information trajectory along a conceptual-to-perceptual gradient, and additionally relays retrieved information to multimodal fronto-parietal brain regions. These findings address the fundamental question of whether memories are more or less complete and truthful reconstructions of past events, or instead are subject to systematic biases that prioritise some types of features over others. Our data suggests that episodic memory retrieval is a dynamic and highly reconstructive process with clear prioritisation of abstract-conceptual over detailed-perceptual information.

neuroscience↗

bioRxiv2022

Phase separation of competing memories along the human hippocampal theta rhythm.

Competition between overlapping memories is considered one of the major causes of forgetting and it is still unknown how the human brain resolves such mnemonic conflict. In the present MEG study, we empirically tested a computational model that leverages an oscillating inhibition algorithm to minimise overlap between memories. We used a proactive interference task, where a reminder word could be associated with either a single image (non-competitive condition) or two competing images, and participants were asked to always recall the most recently learned word-image association. Time-resolved pattern classifiers were trained to detect the reactivated content of target and competitor memories from MEG sensor patterns, and the timing of these neural reactivations was analysed relative to the phase of the dominant hippocampal 3Hz theta oscillation. In line with our preregistered hypotheses, target and competitor reactivations locked to different phases of the hippocampal theta rhythm after several repeated recalls. Participants who behaviourally experienced lower levels of interference also showed larger phase separation between the two overlapping memories. The findings provide evidence that the temporal segregation of memories, orchestrated by slow oscillations, plays a functional role in resolving mnemonic competition by separating and prioritising relevant memories under conditions of high interference.

neuroscience↗

bioRxiv2022

Rhythmic interactions between the mediodorsal thalamus and prefrontal cortex precede human visual perception

The thalamus is much more than a simple sensory relay. High-order thalamic nuclei, such as the mediodorsal thalamus, exert a profound influence over animal cognition. However, given the difficulty of directly recording from the thalamus in humans, next-to-nothing is known about thalamic and thalamocortical contributions to human cognition. To address this, we analysed simultaneously-recorded thalamic iEEG and whole-head MEG in six patients (four female, two male; plus MEG recordings from twelve healthy controls) as they completed a visual detection task. We observed that the phase of both ongoing mediodorsal thalamic and prefrontal low-frequency activity was predictive of perceptual performance. Critically however, mediodorsal thalamic activity mediated prefrontal contributions to perceptual performance. These results suggest that it is thalamocortical interactions, rather than cortical activity alone, that is predictive of upcoming perceptual performance and, more generally, highlights the importance of accounting for the thalamus when theorising about cortical contributions to human cognition.

neuroscience↗

bioRxiv2020

Disentangling the roles of neocortical alpha/beta and hippocampal theta/gamma activity in human episodic memory

To form an episodic memory, we must first process a vast amount of sensory information about a to-be-encoded event and then bind these sensory representations together to form a coherent memory. While these two cognitive capabilities are thought to have two distinct neural origins, with neocortical alpha/beta oscillations supporting information representation and hippocampal theta-gamma phase-amplitude coupling supporting mnemonic binding, evidence for a dissociation between these two neural markers is conspicuously absent. To address this, seventeen human participants completed a sequence-learning task that first involved processing information about three stimuli, and then binding these stimuli together into a coherent memory trace, all the while undergoing MEG recordings. We found that decreases in neocortical alpha/beta power during sequence perception, but not mnemonic binding, correlated with enhanced memory performance. Hippocampal theta/gamma phase-amplitude coupling, however, showed the opposite pattern; increases during mnemonic binding (but not sequence perception) correlated with enhanced memory performance. These results demonstrate that memory-related decreases in neocortical alpha/beta power and memory-related increases in hippocampal theta/gamma phase-amplitude coupling arise at distinct stages of the memory formation process. We speculate that this temporal dissociation reflects a functional dissociation in which neocortical alpha/beta oscillations could support the processing of incoming information relevant to the memory, while hippocampal theta-gamma phase-amplitude coupling could support the binding of this information into a coherent memory trace.

neuroscience↗

bioRxiv2019

Alpha/beta power decreases track the fidelity of stimulus-specific information

Massed synchronised neuronal firing is detrimental to information processing. When networks of task-irrelevant neurons fire in unison, they mask the signal generated by task-critical neurons. On a macroscopic level, mass synchronisation of these neurons can contribute to the ubiquitous alpha/beta (8-30Hz) oscillations. Reductions in the amplitude of these oscillations, therefore, may reflect a boost in the processing of high-fidelity information within the cortex. Here, we test this hypothesis. Twenty-one participants completed an associative memory task while undergoing simultaneous EEG-fMRI recordings. Using representational similarity analysis, we quantified the amount of stimulus-specific information represented within the BOLD signal on every trial. When correlating this metric with concurrently-recorded alpha/beta power, we found a significant negative correlation which indicated that as alpha/beta power decreased, our metric of stimulus-specific information increased. This effect generalised across cognitive tasks, as the negative relationship could be observed during visual perception and episodic memory retrieval. Further analysis revealed that this effect could be better explained by alpha/beta power decreases providing favourable conditions for information processing, rather than directly representing stimulus-specific information. Together, these results indicate that alpha/beta power decreases parametrically track the fidelity of both externally-presented and internally-generated stimulus-specific information represented within the cortex.

neuroscience↗

bioRxiv2018

Hippocampal synchrony and neocortical desynchrony cooperate to encode and retrieve episodic memories

Episodic memories hinge upon our ability to process a wide range of multisensory information and bind this information into a coherent, memorable representation. On a neural level, these two processes are thought to be supported by neocortical alpha/beta desynchronisation and hippocampal theta/gamma synchronisation, respectively. Intuitively, these two processes should couple to successfully create and retrieve episodic memories, yet this hypothesis has not been tested empirically. We address this by analysing human intracranial EEG data recorded during two associative memory tasks. We find that neocortical alpha/beta (8-20Hz) power decreases reliably precede and predict hippocampal \"fast\" gamma (60-80Hz) power increases during episodic memory formation; during episodic memory retrieval however, hippocampal \"slow\" gamma (40-50Hz) power increases reliably precede and predict later neocortical alpha/beta power decreases. We speculate that this coupling reflects the flow of information from neocortex to hippocampus during memory formation, and hippocampal pattern completion inducing information reinstatement in the neocortex during memory retrieval.\n\nSignificance StatementEpisodic memories detail our personally-experienced past. The formation and retrieval of these memories has long been thought to be supported by a division of labour between the neocortex and the hippocampus, where the former processes event-related information and the latter binds this information together. However, it remains unclear how the two regions interact. We uncover directional coupling between these regions, with power decreases in the neocortex that precede and predict power increases in the hippocampus during memory formation. Fascinatingly, this process reverses during memory retrieval, with hippocampal power increases preceding and predicting neocortical power decreases. These results suggest a bidirectional flow of information between the neocortex and hippocampus is fundamental to the formation and retrieval of episodic memories.

neuroscience↗

bioRxiv2017

Data-driven re-referencing of intracranial EEG based on independent component analysis (ICA)

Intracranial recordings from patients implanted with depth electrodes are a valuable source of information in cognitive neuroscience. They allow for the unique opportunity to record brain activity with a high spatial and temporal resolution. To extract the local signal of interest in stereotactic EEG (S-EEG) data, a common pre-processing choice is to re-reference the data with a bipolar montage.\n\nWith bipolar reference, each channel is subtracted from its neighbour in order to reduce commonalities between channels and isolate activity that is spatially confined. We here challenge the assumption that bipolar reference can effectively perform this task. We argue that in order to extract local activity, the distribution of the signal source of interest, as well as the distribution of interfering distant signals and noise sources need to be considered. Those can have a variable spatial extent and are modulated by electrode spacing, location and anatomical characteristics. Those factors are not accounted for by a fixed referencing scheme and bipolar reference can therefore not only decrease the signal to noise ratio (SNR) of the data, but also lead to mislocalization of activity and consequently to misinterpretation of results.\n\nWe promote the perspective of regarding referencing as a spatial filtering operation with fixed coefficients. As an alternative, we propose to use Independent Component Analysis (ICA), to derive filter coefficients that reflect the statistical dependencies of the data at hand. We argue that ICA performs the same task that bipolar referencing pursues, namely undoing the linear superposition of activity and can therefore be used to identify activity that is local. We first describe and demonstrate this procedure on human S-EEG recordings. In a simulation with real data, we then quantitatively show that ICA outperforms the bipolar referencing operation in sensitivity and importantly in specificity when it comes to revealing local time series from the superposition of neighbouring channels.

neuroscience↗