1. Electrophysiology

2. Neurons are Electrical Remember that Neurons have electrically charged membranes they also rapidly discharge and recharge those membranes (graded potentials and action potentials) Review pgs 31 – 50 if this isn’t familiar to you

3. Neurons are Electrical Importantly, we think the electrical signals are fundamental to brain function, so it makes sense that we should try to directly measure these signals –but how?

4. Intracranial and “single” Unit Single or multiple electrodes are inserted into the brain “chronic” implant may be left in place for long periods

5. Intracranial and “single” Unit Single electrodes may pick up action potentials from a single cell An electrode may pick up the signals from several nearby cells –spike-sorting attempts to isolate individual cells

6. Intracranial and “single” Unit Simultaneous recording from several electrodes allows recording of multiple cells

7. Intracranial and “single” Unit Output of unit recordings is often depicted as a “spike train” and measured in spikes/second Stimulus on Spikes

8. Intracranial and “single” Unit Output of unit recordings is often depicted as a “spike train” and measured in spikes/second Spike rate is almost never zero, even without sensory input –in visual cortex this gives rise to “cortical grey” Stimulus on Spikes

9. Intracranial and “single” Unit By carefully associating changes in spike rate with sensory stimuli or cognitive task, one can map the functional circuitry of one or more brain regions

10. Intracranial and “single” Unit Some complications: –Suppose we observe an increase in spike rate in two discrete regions of the brain in response to a sensory stimulus: What are the possible interpretations?

11. Intracranial and “single” Unit Some complications: –Suppose we observe an increase in spike rate in two discrete regions of the brain in response to a sensory stimulus: What are the possible interpretations? 1.Area A “drives” area B 2.Area B “drives” area A 3.Area A and B are controlled by a third area independently

12. Intracranial and “single” Unit Some complications: –Suppose we observe an increase in spike rate in two discrete regions of the brain in response to a sensory stimulus: What are the possible interpretations? 1.Area A “drives” area B 2.Area B “drives” area A 3.Area A and B are controlled by a third area independently and their activity is unrelated How might you differentiate these possibilities

13. Intracranial and “single” Unit How might you differentiate these possibilities Timing of spikes might help: –if A and B are synchronized they are probably functionally related –if A leads B then it is likely to be the first in the signal chain

14. Subdural Grid Intracranial electrodes typically cannot be used in human studies

15. Subdural Grid Intracranial electrodes typically cannot be used in human studies It is possible to record from the cortical surface Subdural grid on surface of Human cortex

16. Electroencephalography It is also possible to record from outside the skull altogether!

17. Electroencephalography pyramidal cells span layers of cortex and have parallel cell bodies their combined extracellular field is small but measurable at the scalp!

18. Electroencephalography The field generated by a patch of cortex can be modeled as a single equivalent dipolar current source with some orientation (assumed to be perpendicular to cortical surface)

19. Electroencephalography Electrical potential is usually measured at many sites on the head surface

20. Electroencephalography Electrical potential is usually measured at many sites on the head surface More is sometimes better

21. Electroencephalography EEG changes with various states and in response to stimuli