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Kriegeskorte, N.

Publications and source records attributed to Kriegeskorte, N..

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Perceived ambiguity of social interactions increases coupling between frontal and temporal nodes of the social brain

Social behaviour is coordinated by a network of brain regions, including those involved in the perception of social stimuli and those involved in complex functions like inferring perceptual and mental states and controlling social interactions. The properties and function of many of these regions in isolation is relatively well understood but little is known about how these regions interact whilst processing dynamic social interactions. To investigate whether social network connectivity is modulated by social context we collected fMRI data from monkeys viewing "affiliative", "aggressive", or "ambiguous" social interactions. We show activation relating to the perception of social interactions along both banks of the superior temporal sulcus, parietal, medial and lateral PFC and caudate nucleus. Within this network we demonstrate that fronto-temporal connectivity are significantly modulated by social context. Crucially, we link the observation of specific behaviours to changes in connectivity within our network. Viewing aggressive or affiliative behaviour was associated with a limited increase in temporo-temporal and premotor-temporal connectivity respectively. By contrast, viewing ambiguous interactions was associated with a pronounced increase in cingulate-cingulate, temporo-temporal, and cingulate-temporal connectivity. We hypothesise that this widespread network synchronisation occurs when cingulate and temporal areas coordinate their activity when more difficult social inferences are made.

neuroscience↗

Rapid event-related, BOLD, NHP: choose two out of three

Human functional magnetic resonance imaging (fMRI) typically employs the blood-oxygen-level-dependent (BOLD) contrast mechanism. In non-human primates (NHP), contrast enhancement is possible using monocrystalline iron-oxide nanoparticles (MION) contrast agent, which has a more temporally extended response function. However, using BOLD fMRI in NHP is desirable for interspecies comparison, and the faster response of the BOLD signal promises to be beneficial to rapid event-related (rER) designs. Here, we used rER BOLD fMRI in macaque monkeys while viewing real-world images, and found visual responses and category-selectivity consistent with previous studies. However, activity estimates were very noisy, suggesting that the lower contrast-to-noise ratio of BOLD, suboptimal behavioural performance, and motion artefacts, in combination, render rER BOLD fMRI challenging in NHP. Previous studies have shown that rER monkey fMRI is possible with MION, despite its prolonged response function. To understand this, we conducted simulations of the BOLD and MION response during rER designs, and found that no matter how fast the design, the greater amplitude of the MION response outweighs the contrast loss caused by greater temporal smoothing. We conclude that although any two of the three elements (rER, BOLD, NHP) have been shown to work well, the combination of all three is particularly challenging.

neuroscience↗

Individual differences among deep neural network models

Deep neural networks (DNNs) excel at visual recognition tasks and are increasingly used as a modelling framework for neural computations in the primate brain. However, each DNN instance, just like each individual brain, has a unique connectivity and representational profile. Here, we investigate individual differences among DNN instances that arise from varying only the random initialization of the network weights. Using representational similarity analysis, we demonstrate that this minimal change in initial conditions prior to training leads to substantial differences in intermediate and higher-level network representations, despite achieving indistinguishable network-level classification performance. We locate the origins of the effects in an under-constrained alignment of category exemplars, rather than a misalignment of category centroids. Furthermore, while network regularization can increase the consistency of learned representations, considerable differences remain. These results suggest that computational neuroscientists working with DNNs should base their inferences on multiple networks instances instead of single off-the-shelf networks.

neuroscience↗

Recurrent networks can recycle neural resources to flexibly trade speed for accuracy in visual recognition

Deep feedforward neural network models of vision dominate in both computational neuroscience and engineering. The primate visual system, by contrast, contains abundant recurrent connections. Recurrent signal flow enables recycling of limited computational resources over time, and so might boost the performance of a physically finite brain or model. Here we show: (1) Recurrent convolutional neural network models outperform feedforward convolutional models matched in their number of parameters in large-scale visual recognition tasks on natural images. (2) Setting a confidence threshold, at which recurrent computations terminate and a decision is made, enables flexible trading of speed for accuracy. At a given confidence threshold, the model expends more time and energy on images that are harder to recognise, without requiring additional parameters for deeper computations. (3) The recurrent models reaction time for an image predicts the human reaction time for the same image better than several parameter-matched and state-of-the-art feedforward models. (4) Across confidence thresholds, the recurrent model emulates the behaviour of feedforward control models in that it achieves the same accuracy at approximately the same computational cost (mean number of floating-point operations). However, the recurrent model can be run longer (higher confidence threshold) and then outperforms parameter-matched feedforward comparison models. These results suggest that recurrent connectivity, a hallmark of biological visual systems, may be essential for understanding the accuracy, flexibility, and dynamics of human visual recognition. Author summaryDeep neural networks provide the best current models of biological vision and achieve the highest performance in computer vision. Inspired by the primate brain, these models transform the image signals through a sequence of stages, leading to recognition. Unlike brains in which outputs of a given computation are fed back into the same computation, these models do not process signals recurrently. The ability to recycle limited neural resources by processing information recurrently could explain the accuracy and flexibility of biological visual systems, which computer vision systems cannot yet match. Here we report that recurrent processing can improve recognition performance compared to similarly complex feedforward networks. Recurrent processing also enabled models to behave more flexibly and trade off speed for accuracy. Like humans, the recurrent network models can compute longer when an object is hard to recognise, which boosts their accuracy. The models recognition times predicted human recognition times for the same images. The performance and flexibility of recurrent neural network models illustrates that modeling biological vision can help us improve computer vision.

neuroscience↗

Rapid invariant encoding of scene layout in human OPA

Successful visual navigation requires a sense of the geometry of the local environment. How do our brains extract this information from retinal images? Here we visually presented scenes with all possible combinations of five scene-bounding elements (left, right and back wall, ceiling, floor) to human subjects during functional magnetic resonance imaging (fMRI) and magnetoencephalography (MEG). The fMRI response patterns in the scene-responsive occipital place area (OPA) reflected scene layout with invariance to changes in surface texture. This result contrasted sharply with the primary visual cortex (V1), which reflected low-level image features of the stimuli, and parahippocampal place area (PPA), which showed better texture than layout decoding. MEG indicated that the texture-invariant scene-layout representation is computed from visual input within ~100 ms, suggesting a rapid computational mechanism. Taken together, these results suggest that the cortical representation underlying our instant sense of the environmental geometry is located in OPA.

neuroscience↗

Analysing linear multivariate pattern transformations in neuroimaging data

Most connectivity metrics in neuroimaging research reduce multivariate activity patterns in regions-of-interests (ROIs) to one dimension, which leads to a loss of information. Importantly, it prevents us from investigating the transformations between patterns in different ROIs. Here, we applied linear estimation theory in order to robustly estimate the linear transformations between multivariate fMRI patterns with a cross-validated Tikhonov regularisation approach. We derived three novel metrics that describe different features of these voxel-by-voxel mappings: goodness-of-fit, sparsity and pattern deformation. The goodness-of-fit describes the degree to which the patterns in an input region can be described as a linear transformation of patterns in an output region. The sparsity metric, which relies on a Monte Carlo procedure, was introduced in order to test whether the transformation mostly consists of one-to-one mappings between voxels in different regions. Furthermore, we defined a metric for pattern deformation, i.e. the degree to which the transformation rotates or rescales the input patterns. As a proof of concept, we applied these metrics to an event-related fMRI data set consisting of four subjects that has been used in previous studies. We focused on the transformations from early visual cortex (EVC) to inferior temporal cortex (ITC), fusiform face area (FFA) and parahippocampal place area (PPA). Our results suggest that the estimated linear mappings are able to explain a significant amount of variance of the three output ROIs. The transformation from EVC to ITC shows the highest goodness-of-fit, and those from EVC to FFA and PPA show the expected preference for faces and places as well as animate and inanimate objects, respectively. The pattern transformations are sparse, but sparsity is lower than would have been expected for one-to-one mappings, thus suggesting the presence of one-to-few voxel mappings. ITC, FFA and PPA patterns are not simple rotations of an EVC pattern, indicating that the corresponding transformations amplify or dampen certain dimensions of the input patterns. While our results are only based on a small number of subjects, they show that our pattern transformation metrics can describe novel aspects of multivariate functional connectivity in neuroimaging data.

neuroscience↗

Faces and voices in the brain: a modality-general person-identity representation in superior temporal sulcus

Face-selective and voice-selective brain regions have been shown to represent face-identity and voice-identity, respectively. Here we investigated whether there are modality-general person-identity representations in the brain that can be driven by either a face or a voice, and that invariantly represent naturalistically varying face and voice tokens of the same identity. According to two distinct models, such representations could exist either in multimodal brain regions (Campanella and Belin, 2007) or in face-selective brain regions via direct coupling between face- and voice-selective regions (von Kriegstein et al., 2005). To test the predictions of these two models, we used fMRI to measure brain activity patterns elicited by the faces and voices of familiar people in multimodal, face-selective and voice-selective brain regions. We used representational similarity analysis (RSA) to compare the representational geometries of face- and voice-elicited person-identities, and to investigate the degree to which pattern discriminants for pairs of identities generalise from one modality to the other. We found no matching geometries for faces and voices in any brain regions. However, we showed crossmodal generalisation of the pattern discriminants in the multimodal right posterior superior temporal sulcus (rpSTS), suggesting a modality-general person-identity representation in this region. Importantly, the rpSTS showed invariant representations of face- and voice-identities, in that discriminants were trained and tested on independent face videos (different viewpoint, lighting, background) and voice recordings (different vocalizations). Our findings support the Multimodal Processing Model, which proposes that face and voice information is integrated in multimodal brain regions.\n\nSignificance statementIt is possible to identify a familiar person either by looking at their face or by listening to their voice. Using fMRI and representational similarity analysis (RSA) we show that the right posterior superior sulcus (rpSTS), a multimodal brain region that responds to both faces and voices, contains representations that can distinguish between familiar people independently of whether we are looking at their face or listening to their voice. Crucially, these representations generalised across different particular face videos and voice recordings. Our findings suggest that identity information from visual and auditory processing systems is combined and integrated in the multimodal rpSTS region.

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GLMdenoise improves multivariate pattern analysis of fMRI data

GLMdenoise is a denoising technique for task-based fMRI. In GLMdenoise, estimates of spatially correlated noise (which may be physiological, instrumental, motion-related, or neural in origin) are derived from the data and incorporated as nuisance regressors in a general linear model (GLM) analysis. We previously showed that GLMdenoise outperforms a variety of other denoising techniques in terms of cross-validation accuracy of GLM estimates (Kay et al., 2013a). However, the practical impact of denoising for experimental studies remains unclear. Here we examine whether and to what extent GLMdenoise improves sensitivity in the context of multivariate pattern analysis of fMRI data. On a large number of participants (31 participants across 4 experiments; 3 T, gradient-echo, spatial resolution 2-3.75 mm, temporal resolution 1.3-2 s, number of conditions 32-75), we perform representational similarity analysis (Kriegeskorte et al., 2008a) as well as pattern classification (Haxby et al., 2001). We find that GLMdenoise substantially improves replicability of representational dissimilarity matrices (RDMs) across independent splits of each participants dataset (average RDM replicability increases from r = 0.46 to r = 0.61). Additionally, we find that GLMdenoise substantially improves pairwise classification accuracy (average classification accuracy increases from 79% correct to 84% correct). We show that GLMdenoise often improves and never degrades performance for individual participants and that GLMdenoise also improves across-participant consistency. We conclude that GLMdenoise is a useful tool that can be routinely used to maximize the amount of information extracted from fMRI activity patterns.

neuroscience↗

From categories to dimensions: spatio-temporal dynamics of the cerebral representations of emotion in voice

Whether the human brain represents emotional stimuli as discrete categories or continuous dimensions is still widely debated. Here we directly contrasted the power of categorical and dimensional models at explaining behavior and cerebral activity in the context of perceived emotion in the voice. We combined functional magnetic resonance imaging (fMRI) and magneto-encephalography (MEG) to measure with high spatiotemporal precision the dynamics of cerebral activity in participants who listened to voice stimuli expressing a range of emotions. The participants also provided a detailed perceptual assessment of the stimuli. By using representational similarity analysis (RSA), we show that the participants perceptual representation of the stimuli was initially dominated by discrete categories and an early (<200ms) cerebral response. These responses showed significant associations between brain activity and the categorical model in the auditory cortex starting as early as 77ms. Furthermore, we observed strong associations between the arousal and valence dimensions and activity in several cortical and subcortical areas at later latencies (>500ms). Our results thus show that both categorical and dimensional models account for patterns of cerebral responses to emotions in voices but with a different timeline and detail as to how these patterns evolve from discrete categories to progressively refined continuous dimensions.\n\nOne Sentence Summary: Emotions expressed in the voice are instantly categorized in cortical processing and their distinct qualities are refined dimensionally only later on.

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Neural dynamics of real-world object vision that guide behaviour

1Vision involves complex neuronal dynamics that link the sensory stream to behaviour. To capture the richness and complexity of the visual world and the behaviour it entails, we used an ecologically valid task with a rich set of real-world object images. We investigated how human brain activity, resolved in space with functional MRI and in time with magnetoencephalography, links the sensory stream to behavioural responses. We found that behaviour-related brain activity emerged rapidly in the ventral visual pathway within 200ms of stimulus onset. The link between stimuli, brain activity, and behaviour could not be accounted for by either category membership or visual features (as provided by an artificial deep neural network model). Our results identify behaviourally-relevant brain activity during object vision, and suggest that object representations guiding behaviour are complex and can neither be explained by visual features or semantic categories alone. Our findings support the view that visual representations in the ventral visual stream need to be understood in terms of their relevance to behaviour, and highlight the importance of complex behavioural assessment for human brain mapping.

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Deep Neural Networks In Computational Neuroscience

The goal of computational neuroscience is to find mechanistic explanations of how the nervous system processes information to give rise to cognitive function and behaviour. At the heart of the field are its models, i.e. mathematical and computational descriptions of the system being studied, which map sensory stimuli to neural responses and/or neural to behavioural responses. These models range from simple to complex. Recently, deep neural networks (DNNs) have come to dominate several domains of artificial intelligence (AI). As the term \"neural network\" suggests, these models are inspired by biological brains. However, current DNNs neglect many details of biological neural networks. These simplifications contribute to their computational efficiency, enabling them to perform complex feats of intelligence, ranging from perceptual (e.g. visual object and auditory speech recognition) to cognitive tasks (e.g. machine translation), and on to motor control (e.g. playing computer games or controlling a robot arm). In addition to their ability to model complex intelligent behaviours, DNNs excel at predicting neural responses to novel sensory stimuli with accuracies well beyond any other currently available model type. DNNs can have millions of parameters, which are required to capture the domain knowledge needed for successful task performance. Contrary to the intuition that this renders them into impenetrable black boxes, the computational properties of the network units are the result of four directly manipulable elements: input statistics, network structure, functional objective, and learning algorithm. With full access to the activity and connectivity of all units, advanced visualization techniques, and analytic tools to map network representations to neural data, DNNs represent a powerful framework for building task-performing models and will drive substantial insights in computational neuroscience.

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Recurrent Convolutional Neural Networks: A Better Model Of Biological Object Recognition Under Occlusion

Feedforward neural networks provide the dominant model of how the brain performs visual object recognition. However, these networks lack the lateral and feedback connections, and the resulting recurrent neuronal dynamics, of the ventral visual pathway in the human and nonhuman primate brain. Here we investigate recurrent convolutional neural networks with bottom-up (B), lateral (L), and top-down (T) connections. Combining these types of connections yields four architectures (B, BT, BL, and BLT), which we systematically test and compare. We hypothesized that recurrent dynamics might improve recognition performance in the challenging scenario of partial occlusion. We introduce two novel occluded object recognition tasks to test the efficacy of the models, digit clutter (where multiple target digits occlude one another) and digit debris (where target digits are occluded by digit fragments). We find that recurrent neural networks outperform feedforward control models (approximately matched in parametric complexity) at recognising objects, both in the absence of occlusion and in all occlusion conditions. Recurrent networks were also found to be more robust to the inclusion of additive Gaussian noise. Recurrent neural networks are better in two respects: (1) they are more neurobiologically realistic than their feedforward counterparts; (2) they are better in terms of their ability to recognise objects, especially under challenging conditions. This work shows that computer vision can benefit from using recurrent convolutional architectures and suggests that the ubiquitous recurrent connections in biological brains are essential for task performance.

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