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fMRI Methods Lecture8 – Electrophysiology & fMRI Neurovascular coupling Iadecola et. al. 2007 Neural activity Ions moving across the membrane “Input” versus “output” of a neuron Energy consumption Brain activity consumes 20% of the body’s energy at rest. Glucose + oxygen => ATP Vasculature replenishes metabolites. Laughlin et. al. Science 2003 Anesthesia reduces brain energy consumption by ~50%. Energy consumption The power required for neural “Signaling” is a sum of both neural output (spikes) and input: Output Input Lennie P. Curr Bio 2003 Neural activity costs Input is more expensive than output. Neurons spend more energy on “listening” than “talking” Dogma: Neural output (firing rate) is the interesting part. Hemodynamic changes Neural input or output? Combination of both? Electrophysiology Different techniques: 1. Intra-cellular recordings 2. Extra-cellular recordings 3. Fluorescence imaging Different spatial resolutions: 1. Single neuron activity 2. Multi unit activity 3. Local field potential Extra-cellular recordings Separate the recorded signal into different components. High frequencies (>500 Hz): Low frequencies (<100 Hz): Simultaneous measurements Measure simultaneous electrophysiology and fMRI and compare. Logothetis et. al. Nature 2001 Simultaneous measurements Before separating electrophysiology into different components Simultaneous measurements LFP and BOLD responses are sustained while MUA seems to adapt very quickly. Anything strange? BOLD – spiking dissociations Several other studies have reported such dissociations… Viswanathan et. al. Nat. Neurosci. 2007 BOLD – spiking dissociations Several other studies have reported such dissociations… Viswanathan et. al. Nat. Neurosci. 2007 Spatial Sampling MUA is a local measure, summing neural spikes only of neurons surrounding the immediate electrode tip. LFP and BOLD are wider measures, summing dendritic/synaptic activity several mm surrounding the electrode. Sampling bias Multi unit activity is mainly generated by large layer 5 pyramidal cells. These are the main “output” neurons of the cortex. LFP and BOLD sum across all cell sizes in all layers. Cortical structure In cortex, 80% of a neurons output synapses are located within 1 mm of its soma. Strong recurrent innervation. Only 6% of V1 synapses (mostly layer 4) are from thalamic neurons. Input without output? Increased LFP without spiking? BOLD, LFP, and spikes Epilepsy patients implanted with electrodes in auditory cortex: Mukamel et. al. Science 2005 BOLD, LFP, and spikes Neural activity correlated with fMRI: BOLD, LFP, and spikes Different LFP frequencies showed different relationship to BOLD: Optogenetics Hyung Lee et. al. Nature 2010 Optogenetics Inject virus into motor cortex. Axons of infected cells reach thalamus. Stimulate in motor cortex and measure activity in both locations. During rest What about spontaneous activity? Shmuel et. al. HBM 2008 During rest Significant correlations between neural activity and BOLD during rest… Negative BOLD? Shmuel et. al. Nat. Neurosci. 2006 Negative BOLD? Negative BOLD? Cerebellum Subcortical brain areas might demand more caution. Architecture is different: no recurrent innervation. GABA agonist halts PC spikes BOLD continues There is a difference between input and output. BOLD coupled to input. Caeser et. al. PNAS 2003 One more thought The effects of neuro-modulators (caffeine, hormones, noradrenalin, dopamine, serotonin, etc…) on particulars of neural activity and neurovascular coupling are unknown. To the lab