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The brain at rest
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Spontaneous rhythms in a dish
Connected neural populations tend to synchronize and oscillate together.
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EEG Alpha (10 Hz)
Over occipital electrodes.
Different time scales of synchronization
Close eyes
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Slow changes
Ultra slow changes in neural firing rates and LFP in auditory cortex of a resting and sleeping human.
Nir Y., Nat. Neurosci. (2010)
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Slow changes
Similar fluctuations in fMRI signal during rest
Nir Y., Neuroimage (2006)
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Energy costs
Brain takes up 20% of the metabolites in the body during rest.
Nir Y., Neuroimage (2006)
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Energy costs
Energy cost breakdown for running the cortex.
Most of the energy is spent on neural signaling.
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Energy costs
Energy consumption measured with PET (radioactive glucose consumption) during 4 conditions.
Which is rest and which is task?
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Energy costs
Brain is always at max power: similar costs in all conditions
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Information encodingGiven that energy consumption is almost constant:
Only relatively small populations of neurons respond strongly at any given time.
Relatively local, transient changes in neural activity represent stimuli evoked responses.
Hence the importance of: Attention and neurovascular coupling
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Structure of rest activity
Spontaneous neural activity during rest and sleep is not random “noise”.
Large neural populations are synchronized.
What are the characteristics of this activity?
How can we study them? No experimental structure?
Are there differences between rest, sleep, anesthesia?
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Local spatial structure
Anesthetized cat primary visual cortex
Orientation columnar map – voltage sensitive dye imaging
Kenet, Nature (2003)
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Local spatial structure
Many occurrences of strong correlation between spontaneous and evoked columnar maps.
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Local spatial structure
Correlation between spontaneous “snapshots” and particular orientation columnar maps.
Visual cortex randomly moves from one map to another in the absence of stimulation (relatively quick transitions).
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Large spatial structure
Cortical areas with similar functionality (e.g. right and left auditory cortex) show strong and selective correlations.
Very slow hemodynamic changes over time.
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Functional connectivityAreas that are connected anatomically because of shared
functionality will be active together.
In reality this is just correlation – problematic interpretation
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Inter-hemispheric correlations
Strongest correlations are between corresponding locations in the two hemispheres
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Default mode system
Three areas that show reduced activity during “external” tasks (e.g. visual, auditory, somatosensory stimulation)
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Default mode system at rest
Is the brain separated into two general antagonistic networks?
Measuring default mode system correlations during rest.
“Intrinsic” ROI“Extrinsic”
ROI
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Source of fMRI correlations
Several “less interesting “ sources contribute to an fMRI signal. Are they driving correlations during rest?
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Source of fMRI correlationsSimultaneous fMRI and electrophysiology:
Shmuel et. al. HBM 2008
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Source of fMRI correlations
Significant correlations between neural activity and BOLD during rest…
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Source of fMRI correlations
Similar inter-hemispheric correlations in epilepsy patients
Nir Y., Nat. Neurosci. (2010)
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Source of fMRI correlations
Correlations during rest correspond to anatomically connected areas that are commonly active during task.
Fox, Nat. Rev. Neurosci. (2007)
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Patient lacking corpus callosum
Inter-hemisphericcorrelations disappear after section of corpus callosum.
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Spontaneous and evoked activityThere’s a lot of variability in the neural response to the same repeating visual stimulus.
Arieli, Science (1996)
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Spontaneous and evoked activity
The spontaneous state right before stimulation predicted the amplitude of the response 42 ms later.
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Spontaneous and evoked activity
Changing the spatial response pattern in a predictable way.
Is the evoked response to atrial the sum ofspontaneous and evokedactivity?
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Motor response variability
Fox, Nat. Neurosci. (2006)
Right hand button press LH ROI
Functionally correlated RH ROI
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Motor response variability
LH responses RH responses
Subtracting out RH spontaneous activity, allowed reduction of trial by trial evoked response variability.
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Diagnosis by rest scans
If resting state correlations represent the functionality of the brain, abnormal correlations may represent abnormal functionality and enable an easy form of diagnosis.
Build a distribution of “normal” correlation values and test whether these deviate in particular neurological or psychiatric disorders.
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Alzheimer’s disease
Weaker resting state correlations between “default mode” network (precuneus area) and hippocampi.
Fare comparison?
Control Alheimer’s
Greicius, PNAS (2004)
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Alzheimer’s disease
Direct comparison:
Voxels that showed stronger correlation with PPC timecourse in controls versus Alzheimer’s patients.
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Comma
Patients in reduced states of awareness show decreased inter-hemispheric correlations.
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Autism
1-4 year old children with autism or language delay, during natural sleep.
Strong inter-hemispheric correlations are already evident at extremely young ages.
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Autism
Decreased inter-hemispheric correlations in language areas
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Autism
Inter-hemispheric correlation strength predicts language ability and autism symptom severity.
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Summary
Spontaneous brain activity is the largest and least understood component of brain function.
Different temporal and spatial scales of organization.
Immense importance – development, maintenance, health.
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To the lab!