Source: Laboratories of Jonas T. Kaplan and Sarah I. Gimbel—University of Southern California
Imagine the sound of a bell ringing. What is happening in the brain when we conjure up a sound like this in the "mind's ear?" There is growing evidence that the brain uses the same mechanisms for imagination that it uses for perception.1 For example, when imagining visual images, the visual cortex becomes activated, and when imagining sounds, the auditory cortex is engaged. However, to what extent are these activations of sensory cortices specific to the content of our imaginations?
One technique that can help to answer this question is multivoxel pattern analysis (MPVA), in which functional brain images are analyzed using machine-learning techniques.2-3 In an MPVA experiment, we train a machine-learning algorithm to distinguish among the various patterns of activity evoked by different stimuli. For example, we might ask if imagining the sound of a bell produces different patterns of activity in auditory cortex compared with imagining the sound of a chainsaw, or the sound of a violin. If our classifier learns to tell apart the brain activity patterns produced by these three stimuli, then we can conclude that the auditory cortex is activated in a distinct way by each stimulus. One way to think of this kind of experiment is that instead of asking a question simply about the activity of a brain region, we ask a question about the information content of that region.
In this experiment, based on Meyer et al., 2010,4 we will cue participants to imagine several sounds by presenting them with silent videos that are likely to evoke auditory imagery. Since we are interested in measuring the subtle patterns evoked by imagination in auditory cortex, it is preferable if the stimuli are presented in complete silence, without interference from the loud noises made by the fMRI scanner. To achieve this, we will use a special kind of functional MRI sequence known as sparse temporal sampling. In this approach, a single fMRI volume is acquired 4-5 s after each stimulus, timed to capture the peak of the hemodynamic response.
1. Participant recruitment
2. Pre-scan procedures
3. Provide instructions for the participant.
4. Put the participant in the scanner.
5. Data collection
6. Data analysis
Figure 1: Region of interest tracing. The surface of the planum temporale has been traced on this participant's high-resolution anatomical image, and is shown here in blue. In green is the control mask of the frontal pole. These voxels will be used for MVPA analysis.
The average classifier accuracy in the planum temporale across all 20 participants was 59%. According to the Wilcoxon Signed-Rank test, this is significantly different from chance level of 33%. The mean performance in the frontal pole mask was 32.5%, which is not greater than chance (Figure 2).
Figure 2. Classification performance in each participant. For three-way classification, chance performance is 33%. According to a permutation test, the alpha level of p < 0.05 corresponds to 42%.
The permutation test found that only 5% of the permutations achieved accuracy greater than 42%; thus, our statistical threshold for individual subjects is 42%. Nineteen of the 20 subjects had classifier performance significantly greater than chance using voxels from the planum temporale, while none had performance greater than chance using voxels from the frontal pole.
Thus, we are able to successfully predict from patterns of activity in auditory cortex which of the three sounds the participant was imagining. We were not able to make this prediction based on activity patterns from the frontal pole, suggesting that the information is not global throughout the brain.
MVPA is a useful tool for understanding how the brain represents information. Instead of considering the time-course of each voxel separately as in a traditional activation analysis, this technique considers patterns across many voxels at once, offering increased sensitivity compared with univariate techniques. Often a multivariate analysis uncovers differences where a univariate technique is not able to. In this case, we learned something about the mechanisms of mental imagery by probing the information content in a specific area of the brain, the auditory cortex. The content-specific nature of these activation patterns would be difficult to test with univariate approaches.
There are additional benefits that come from direction of inference in this kind of analysis. In MVPA we start with patterns of brain activity and attempt to infer something about the mental state of the participant. This kind of "brain-reading" approach can lead to the development of brain-computer interfaces, and may allow new opportunities for communication with those with impaired speech or movement.
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