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Advance fMRI analysis: MVPA
Sara Fabbri Brain and Mind Institute Western University Advance fMRI analysis: MVPA
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Overview Why MVPA? How MVPA “Mind-reading” MVPA classifier
MVPA correlation Representational similarity approach “Mind-reading”
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Response of a neuron from the human medial temporal lobe
Picture number Picture Raster plots for various trials (from top to bottom) Firing rate onset offset Quiroga et al., 2005, Nature
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Response of a neuron from the human medial temporal lobe
Quiroga et al., 2005, Nature
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Response of a neuron from the human medial temporal lobe
Selective response to Jennifer Aniston pictures Neuron 1 “likes” Jennifer Aniston Quiroga et al., 2005, Nature
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Firing Rate Firing Rate Firing Rate Neuron 1 “likes” Jennifer Aniston
Julia Roberts Neuron 3 “likes” Brad Pitt Firing Rate Firing Rate Firing Rate
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fMRI spatial resolution: 1 voxel
high activity fMRI spatial resolution: 1 voxel 3 mm Fusiform Face Area (FFA) 3 mm 3 mm 3 mm low activity 3 mm A voxel might contain millions of neurons, so the fMRI signal represents the population activity
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Firing Rate Firing Rate Firing Rate Activation
Neuron 1 “likes” Jennifer Aniston Neuron 2 “likes” Julia Roberts Neuron 3 “likes” Brad Pitt Even though there are neurons tuned to each actor, the population as a whole shows no preference Firing Rate Firing Rate Firing Rate Activation
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Limitation of Subtraction Logic
Are there neurons that prefer Jennifer to Julia in the voxel? > Subtraction Activation Activation No preference
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Two Techniques with “Subvoxel Resolution”
“subvoxel resolution” = the ability to investigate coding in neuronal populations smaller than the voxel size being sampled Multi-Voxel Pattern Analysis (MVPA or decoding or “mind reading”) fMR Adaptation (or repetition suppression or priming)
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Multi-Voxel Pattern Analyses
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fMRI spatial resolution: 1 voxel
high activity fMRI spatial resolution: 1 voxel 3 mm low activity 3 mm
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Region Of Interest (ROI): group of voxels
high activity 3 mm 3 mm low activity 3 mm
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Standard fMRI Analysis
FACES HOUSES trial 1 trial 1 trial 2 trial 2 trial 3 trial 3 Average Summed Activation
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4 mm Full-width-Half-Maximum (FWHM)
Spatial Smoothing 4 mm Full-width-Half-Maximum (FWHM) 7 mm FWHM 10 mm FWHM No smoothing Application of a Gaussian low-pass spatial filter to fMRI data resulting in spreading the activation across adjacent voxels Why? increase signal-to-noise facilitate intersubject averaging
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Effect of Spatial Smoothing: reduce spatial resolution
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Why Multi-Voxel Pattern Analysis (MVPA)?
Patterns carry more information than the average across a pattern Average Pattern Standard fMRI Analysis MVPA fMRI Analysis
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We have seen that the pattern of activity might be informative but it is lost in the standard fMRI analysis Questions?
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How Multi-Voxel Pattern Analysis?
MVPA classifier MVPA correlation Representational similarity approach
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1. MVPA CLASSIFIER
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FACES HOUSES trial 1 trial 1 Training Trials trial 2 trial 2 trial 3
… … Can an algorithm correctly “guess” trial identity better than chance (50%)? Test Trials (not in training set)
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Voxel 1 Voxel 2 Activity in Voxel 1 Activity in Voxel 2 Faces Houses
Each dot is one measurement (trial) from one condition (red circles) or the other (green circles) Activity in Voxel 2 Faces Houses
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Training set Test set Activity in Voxel 1 Activity in Voxel 2 Faces Houses Classifier
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THIS STEP IS THE CORE OF THE ANALYSIS BECAUSE IT TESTS THE ABILITY OF THE CLASSIFIER TO GENERALIZE KNOWLEDGE TO NEW DATA Test set Activity in Voxel 1 Activity in Voxel 2 Faces Correct 6 Classifier Accuracy = = = 75 % Houses 8 Incorrect Classifier
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Faces Houses In a, the classifier can operate on single voxels because the response distributions are separable within individual voxels. Cox and Savoy (2003)
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Also standard fMRI analysis can detect this difference
Faces Houses Also standard fMRI analysis can detect this difference Cox and Savoy (2003)
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Faces Houses In b, the classifier cannot operate on single voxels because the response distributions are overlapping within individual voxels. Cox and Savoy (2003)
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Standard fMRI analysis cannot detect this difference
Faces Houses Standard fMRI analysis cannot detect this difference Cox and Savoy (2003)
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In c, the non-linear classifier draws decision boundaries other than straight lines
Cox and Savoy (2003)
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Standard fMRI analysis cannot detect this difference
Cox and Savoy (2003)
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Support Vector Machine (SVM)
SVM is a linear decision boundary that discriminates between two sets of points with the constrain of a greater distance from the closest points on both sides. The response patterns closest to the decision boundary (yellow circles) that defined the margins are called “support vectors”. Mur et al., 2009 Mur et al., 2009
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Choices to make Feature selection to remove noisy and/or uninformative voxels before classification: Limit the analysis to specific anatomical regions Compute univariate (voxel-wise) statistics Cross-validation options: Leave-one-trial out: for each condition train the classifier with all but one trial and use the left out subset as test data. Repeat this procedure until each subset has been used as test data once. Leave-one-run out: train the classifier with all but one runs and test on the excluded run. Repeat until all runs are used as test data once. Performance evaluation options: T-test versus 50% chance decoding Permutation test: train and test the classifier with labels (e.g. faces and houses) randomly assigned to the data. Compare classifier performance on real data with performance on randomized data.
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Example of MVPA classifier approach: decoding future actions
Gallivan et al., 2011
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Conditions Gallivan et al., 2011
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Delayed-paradigm Gallivan et al., 2011
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PREDICT FUTURE EFFECTOR
PREDICT THE UPCOMING REACH OR SACCADE Gallivan et al., 2011
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PREDICT FUTURE DIRECTION
PREDICT THE UPCOMING MOVEMENT DIRECTION Gallivan et al., 2011
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PREDICT FUTURE ACTION! Gallivan et al., 2011
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What else can we examine?
So far we’ve only examined the question: Can we discriminate Condition 1 vs. Condition 2 from brain activation patterns? New question: Are different stimuli similarly represented in certain ROIs? Answer: Cross-decoding: train to discriminate between Condition 1 and 2 and test classifier accuracy with Condition 3 and 4 Fraser Smith & Jason Gallivan
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Cross-trial-type decoding
Training set Activity in Voxel 1 Activity in Voxel 2 Faces Houses Classifier Fraser Smith & Jason Gallivan
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Cross-trial-type decoding
Test set Activity in Voxel 1 Activity in Voxel 2 Animals Tools Classifier Fraser Smith & Jason Gallivan
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Similar spatial encoding across the eye and the hand
Train to discriminate movement directions for one effector (HandL vs HandR) and test it on the other effector (EyeL vs EyeR) Gallivan et al., 2011
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Effector specificity across different movement directions
Train to discriminate the effector for one movement directions (EyeL vs HandL) and test it on the other direction (EyeR vs HandR) Gallivan et al., 2011
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Effector specificity across different movement direction
Gallivan et al., 2011
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We have seen how MVPA classifier can be used to distinguish between different stimulus categories based on the pattern of activity Questions?
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2. CORRELATION APPRAOCH
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MVPA correlation approach
Faces Houses trial 1 trial 1 trial 1 trial 2 trial 2 trial 2 trial 3 trial 3 trial 3 trial 3 Average Summed Activation
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MVPA correlation approach
Faces Houses trial 1 trial 1 trial 1 trial 1 trial 2 trial 2 trial 2 trial 2 trial 3 trial 3 trial 3 trial 3 trial 3 Average Summed Activation The same category evokes similar patterns of activity across trials
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MVPA correlation approach
Faces Houses trial 1 trial 1 trial 1 trial 2 trial 2 trial 2 trial 3 trial 3 trial 3 trial 3 Average Summed Activation Similarity Within the same category
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MVPA correlation approach
Faces Houses trial 1 trial 1 trial 1 trial 1 trial 2 trial 2 trial 2 trial 2 trial 3 trial 3 trial 3 trial 3 Average Summed Activation Similarity Between different categories
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The brain area contains distinct information about faces and houses
Within-category similarity Between-category similarity > The brain area contains distinct information about faces and houses
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Category-specificity of patterns of response in the ventral temporal cortex
Haxby et al., 2001, Science
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Within-category similarity
Category-specificity of patterns of response in the ventral temporal cortex SIMILARITY MATRIX ODD RUNS high EVEN RUNS similarity low Within-category similarity Haxby et al., 2001, Science
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Between-category similarity
Category-specificity of patterns of response in the ventral temporal cortex SIMILARITY MATRIX ODD RUNS high EVEN RUNS similarity low Between-category similarity Haxby et al., 2001, Science
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fMRI activity high similarity low Bracci et al., 2011
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Similar representation for tools and hand not identified from the standard univariate analysis
fMRI activity high similarity low Bracci et al., 2011
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Searchlight analysis We have seen how to search for patterns of information within a region of interest, but what if we want to know “Where in the brain does the activity pattern contain information about the experimental condition?” Scanned the brain with a spherical multivariate “searchlight” centered on each voxel in turn (optimal searchlight has radius of 4 mm, that contains 33 2-mm-isovoxel voxels) Compute multivariate effect statistic at each location Kriegeskorte et al., 2006
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We have seen how MVPA correlation can measure similarities and differences between patterns of activity across conditions Questions?
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3. REPRESENTATIONAL SIMILARITY APPROACH (RSA)
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PRE-DEFINED CATEGORIES
CATEGORY A CATEGORY B Candies in different boxes are different because they have different prices
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Which dimension to choose for the classification?
HOW ARE THESE OBJECTS CLASSIFIED IN THE BRAIN (BY PRIC€, COLOR, SHAPE, FLAVOUR OR OTHER DIMENSIONS)?
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Representational similarity approach (RSA)
Differently from the MVPA correlation, RSA does not separate stimuli into a-priori categories MVPA correlation RSA high low similarity ODD RUNS EVEN RUNS Kriegeskorte et al (2008)
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No class boundaries! C1 high . . CONDITIONS TRIALS similarity low C96
Fraser Smith & Jason Gallivan
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TEST VARIOUS PREDICTIONS
high low similarity Kriegeskorte et al (2008)
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TEST VARIOUS PREDICTIONS
Which prediction matrix is more similar to the real data? high low similarity REAL DATA Kriegeskorte et al (2008)
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Interspecies comparisons
Man + monkey?? similarity high low Kriegeskorte et al., 2008 Fraser Smith & Jason Gallivan
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Activation- vs. information-bases analysis
Activation-based (standard fMRI analysis): regions more strongly active during face than house perception Information-based (searchlight MVPA analysis): regions whose activity pattern distinguished the two categories 35 % of voxels are marked only in the information-based map: category information is lost when data are smoothed Kriegeskorte, Goebel & Bandettini, 2006
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Activation- vs. information-based analysis
Mur et al., 2009, Social Cognitive and Affective Neuroscience
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Limitations of MVPA Good classification indicates presence of information (no necessarily neuronal selectivity) (Logothetis, 2008). E.g. successful face decoding in primary visual cortex Pattern-classifier analysis requires many decisions that affect the results (see Misaki et al., 2010)
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We have seen how RSA can be used for exploratory analysis with complex stimulus set
To summarize, the three MVPA methods are powerful especially to study distributed representations Questions?
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“Mind-Reading”: Reconstructing new stimuli from brain activity
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Reconstruct new images
The model is trained to classify the pattern on brain activity in primary visual cortex during observation of several hundred random images The model can accurately reconstruct arbitrary images, including geometric shapes on a single 2 sec volume without any prior information about the image Miyawaki et al., 2008
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Reconstruct new movies
3 participants watched hours of movie trailers during fMRI Reconstruction of new movies from brain activity during similar clips from YouTube Presented movies Reconstructed movies (Nishimoto et al., 2011
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Lie detector Non-linear classifier applied to fMRI data to discriminate spatial patterns of activity associated to lie and truth in 22 individual participants. 88% accuracy to detect lies in participants not included in the training (Davatzikos et al., 2005)
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Lie detector Non-linear classifier applied to fMRI data to discriminate spatial patterns of activity associated to lie and truth in 22 individual participants. 88% accuracy to detect lies in participants not included in the training The real world is more complex!
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Reconstruct dreams Measure brain activity while 3 participants were asleep and ask them to describe their dream when awake Comparison between brain activity during sleep and vision of pictures of categories frequently dreamt Activity in higher order visual areas (i.e. FFA) could successfully (accuracy of 75-80%) decode the dream contents 9 seconds before waking the participant! Abstract SfN 2012: Dreaming is a subjective experience during sleep, often accompanied by visual contents, whose neural basis remains unknown. Previous dream research attempted to link physiological states with dreaming, but did not demonstrate how the specific contents of visual experiences during dreaming are represented in the brain activity patterns. The recent advent of neural decoding has allowed for the decoding of various contents of visual experience from brain activity patterns. The technique can thus be used to examine the neural representation of dreams by testing whether neural decoders can predict dream contents from brain activity patterns. Here we performed decoding analyses on semantically labeled human fMRI signals measured from three male subjects during dreaming. To collect dream data efficiently, we measured fMRI signals and collected reports about subjective experiences during hypnagogic periods. We developed a multiple-awakening procedure, in which subjects were awakened when a specific EEG pattern was observed, were asked to freely describe their visual experiences just before awakening, and were then asked to sleep again. We repeated this procedure until we collected over 200 reports in a total of hours of experiment time for each subject. Multiple “synsets,” synonym sets defined in the English “WordNet” lexical database, that correspond to words describing reported visual contents were used to label averaged fMRI volumes during a 9 s period before each awakening. We first performed pairwise classification analyses for all pairs of synsets using fMRI signals in the early (V1-V3) and the higher (around LOC, FFA, and PPA) visual cortices during dreaming (dream-trained decoder). The decoding performance showed a distribution that was significantly higher than chance level in the higher visual areas. We next examined whether “stimulus-trained decoders” that were trained with fMRI signals evoked by natural image viewing could decode the dream contents. Results showed that the stimulus-trained decoders successfully predicted the dream contents more accurately in the higher visual cortex than in the early visual cortex. These results demonstrate that fMRI signals in the visual cortex, especially in the higher visual areas, represent specific visual contents of dreams, allowing for the prediction of dream contents. Furthermore, it supports the hypothesis that dreaming and perception may share neural representations in the higher visual areas. Horikawa, Tamaki, Miyawaki and Kamitani, Society for Neuroscience 2012
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Thousand of categories in the real world
Which of the thousand of categories are represented similarly in the brain and which aren’t? Measure brain activity from 5 subjects while watching movies See example movies:
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Shared Semantic Space from brain activity during observation of movies
Similar colors for categories similarly represented in the brain Huth et al., 2012
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Shared Semantic Space from brain activity during observation of movies
Similar colors for categories similarly represented in the brain People and communication verbs are represented similarly Huth et al., 2012
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Continuous Semantic Space across the surface
Each voxel is colored accordingly to which part of the semantic space is selective for
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Continuous Semantic Space across the surface
Click on each voxel to see which categories it represents FUSIFORM FACE AREA
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Continuous Semantic Space across the surface
Click on each category to see how it is represented in the brain FACE
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The possibility to decode brain activity has great potentials for brain-computer applications
Let’s watch a video!
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The end
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