January 19, 2009

Phase Alignment of Neocortical Gamma Oscillations by Hippocampal Theta Waves

An empty brain is the devil’s workshop, goes the proverb. Actually, the brain is never empty. Even in our deepest slumber, the brain continues to weave waves of electrical rhythms that can be seen with the aid of electroencephalogram or EEG. When we place electrodes on the scalp or on the cortex (inside the skull), and amplify the faint signals via bioinstrumentation amplifier, we can lay our hands on these fluctuating rhythms. (More on the electronics of EEG may be found at the OpenEEG project site).

We have as many as 100 billion neurons in the brain. In the superficial layers of the cortex, the neurons have numerous dendrites branching out from the soma or cell body (shown in grey oval in this picture).diagrammatic representation of cortical dipole with dendritic treesThese neurons have been compared to a forest of trees where the branches are the dendrites and the trunk the axon. These dendrites make extensive connections among each other. They also get connections from the axon collaterals of neighboring axons (i.e. the 'trunks' of other trees connect to these 'twigs' by offshoot from the trunks). Since there are a lot of axons converging on the dendrites of each neuron, and given the fact that these axons can be excitatory (red) or inhibitory (green) depending on the neurotransmitter, the sum of input may be either negative or positive (with respect to the cell body). Thus an alternating current (cortical dipole) will flow between the shifting dendrites and the soma. This along with thalamocortical oscillations produces the EEG waves.

The brain doesn’t churn out the rhythm just like that. Had the neurons fired randomly the oscillations would have cancelled out.EEG showing alpha, beta and other brainwavesEEG waves occur due to synchronous discharge of neurons producing the alpha, beta, theta, gamma and other telltale waves. Like all other electrical waves, they too have a frequency and amplitude. Alpha waves, for example, have a frequency of 8-12 Hz (cycles per second) and an amplitude ranging from 50-100 microvolt when recorded from the scalp, and it is found when a person is resting comfortably with eyes closed and the mind wandering. On the other hand, gamma rhythm has a frequency of 30-80 Hz, and it is found when a person is deeply engrossed on some work.

It was known for a long time that the hippocampus exerted a role in learning by fostering long term potentiation (LTP) by aligning the neocortex, where memories are stored. The mechanisms behind this are now emerging. Sirota et al and Siapas et al have analyzed rat brains and found out that there were many localized gamma oscillators within the brain that gave rise to neocortical gamma bursts. These oscillators had varying frequencies but they phase aligned themselves with the arrival of hippocampal theta waves. A large fraction of pyramidal cells and interneurons too were phase aligned to the hippocampal theta rhythm.Bar magnet showing lines of forceThis is similar to a bar magnet aligning iron dust or other ferromagnetic materials by virtue of its magnetic field. Apart from the cerebral cortex, the cerebellar cortex and the hippocampus too can generate brain waves. Such a mechanism may explain the orchestration of many parts of the cortex (and hence the memory engrams they contain); and data synchronization and downloading to the hippocampus for memory retrieval. It also shows how hippocampus does the ‘indexing’ of cortical contents. These experiments throw light on neuronal plasticity and information flow, and may be someday they could help clinicians in fighting memory loss as it occurs in neurodegenerative diseases like Alzheimer’s disease.

Last modified: never
Prefrontal Phase Locking to Hippocampal Theta Oscillations
Athanassios G. Siapas, Evgueniy V. Lubenov and Matthew A. Wilson. doi:10.1016/j.neuron.2005.02.028
ResearchBlogging.orgA SIROTA, S MONTGOMERY, S FUJISAWA, Y ISOMURA, M ZUGARO, G BUZSAKI (2008). Entrainment of Neocortical Neurons and Gamma Oscillations by the Hippocampal Theta Rhythm Neuron, 60 (4), 683-697 DOI: 10.1016/j.neuron.2008.09.014


The Spiral Mindscope said...

I thought the generation of electrical activity occurred down in peri-aqueductal grey area in the brain-stem, not the cortex. That's the bit that statutory hanging damages irrevocably and that kills the person. Cortex merely outputs electrical activity - that's why one could survive despite being partially decorticated.

I liked the article and especially the analogy with magnetic field. What is the human brain's electrical field beyond the waves? and how will it behave within a stronger field?

amiya said...

Jay, I like your question. Judicial hanging kills by asphyxia, apoplexy or more commonly a combination of both, apart from other causes such as fracture/dislocation of cervical vertebrae and pressure on the 'vital centers' in the medulla.

While decortication is compatible with life in humans, it causes rigidity (spasticity) and the person is not conscious. For consciousness gamma oscillations are required (http://physiology-physics.blogspot.com/2007/06/consciousness-inside-story.html)

Electrical field beyond the waves is a fantastic horizon that needs to be explored. I look forward to see the answer in YOUR blog -:)One possibility (and now a reality)is thought controlled devices (http://physiology-physics.blogspot.com/2006/12/telepathy-nebulous-domain-of-mystique.html)

The Spiral Mindscope said...

HI Amiya

I was referring in my response to the comment that EEG waves are 'created' in the neocortex. EEG waves are the output of an electrical 'generator' that resides in the brain-stem. You're right of course that for someone to be 'conscious', neorcortical EEG activity is a must. That's of course true for higher mammals only, not for lower or licencephalic animals, who dont have neocortex and subsist on paleo-cortex alone. They are conscious too but only in an intuitive way, cannot recall much of what they experience except those learnt through classical and operant conditioning.

Finally, gamma-waves represent what is being termed the 'binding' factor; i.e. it allows multiple representations and rapid permutations and computations of an object of human attention. Onecan still be 'conscious' without much gamma- wave oscillations. What they will lack though is a 'conscious' and detailedmemory of those 'conscious' awareness after the stimulus is over.

I loved the link - and of course I shall post something related on my blog and provide a link to a most stimulating article.

amiya said...

While science have established that 'delta waves' originated from within the cortex, doubts exist regarding 'alpha rhythm'. Besides the 'reticular nuclei' around the thalamus and intrathalamic nuclei, medulla perhaps play a role too. There's even this view that alpha waves originate from the heart! To keep the blog article short, I didn't include them. Moreover, typing 'em all is quite a boring job. Thanks for pointing.

Shaheen Lakhan said...

Thanks for submitting this post to our blog carnival. We just published the 43rd edition of Brain Blogging and your article was featured!

Thank you.


Anonymous said...

It is rather interesting for me to read the post. Thank you for it. I like such themes and anything connected to this matter. I definitely want to read more soon.