Nicholas Haley 

12/9/2021 
haleynp@mail.uc.edu 
M11960517 
 
Cellular Mechanisms Underlying Brain Waves 

Brain waves, or neural oscillations, are characterized by repetitive synchronous firing of

neuron pools. These waves take place in various regions of the brain and are generally associated

with mental states or functions. They occur at all times and appear differently in states ranging

from engaged problem solving to deep sleep.  

Brain waves are categorized by frequency. These frequencies range from 0.5 Hz all the

way to 100 Hz. The major categories include delta (0.5-3 Hz), theta (3.5-7 Hz), alpha (8-3 Hz),

beta (13-30 Hz), and gamma (30-100 Hz) (Lee). The frequency ranges are delineated by speed

and different sources provide different frequency ranges for each of the categories. These

categories are further divided into subtypes. These frequency categories are often associated with

mental states. For example, beta waves are commonly associated with an alert state of mind

while theta waves are found during deep sleep.  

Low frequency brain waves are associated with low levels of consciousness while high

frequency brain waves are associated with an alert consciousness (Scientific American). While

this is generally true, multiple brain waves usually take place in the brain at the same time. What

this is really referring to is the dominant waveform, or the wave frequency that appears the most

consistently during a particular mental state.  

The frequency that has been most heavily studied is the gamma frequency. One reason

for studying it is that gamma waves are relatively easy to induce. Gamma waves are produced in

the prefrontal cortex during mental activity that requires focus. One way to induce these brain

waves is by having subjects perform focused meditation, such as simply focusing on their
breathing, while performing and EEG to measure the brain waves. Interestingly, one study found

that the duration of gamma waves seems to correlate with the duration of focus (Goleman).

People who are classified as being high level meditators are able to sustain gamma frequencies

much longer, and some of the even more advanced meditators produce gamma waves even when

not performing meditation (Goleman). This suggests that brainwaves are malleable and further

validates the idea that brain waves are associated with mental states.   

Brain waves are generated by repetitive excitation and inhibition of neurons in a neuron

pool (Buzsáki). These collective firings induce electromagnetic fields in accordance with the

collective changes in membrane potential brought about by action potentials. What is truly

incredible is that all these action potentials in a neuron pool can coincide to create an oscillating

electromagnetic field.  

These brain waves can be detected and studied using EEGs and MEGs. EEGs and MEGs

allow scientists to differentiate what frequencies of brain waves are taking place in the brain.

These EEG readings can appear rather chaotic because they show multiple frequencies at once.

Through computer software, its possible to use spectral analysis to pull out specific frequencies

and determine where individual frequencies originate in the brain during a particular mental state

(Cole). 

Although EEG and MEG are able to detect brainwaves quite reliably and with high

resolution, they are limited to being able to detect brainwaves from more superficial regions of

the brain. Another limitation in studying brainwaves is being able to consistently induce the

brain waves that are desired to study. For example, low frequency waves, like theta waves, occur

during relaxed states and deep sleep. Being able to get subjects to fall into a deep sleep in a lab
setting is likely to be quite challenging. Therefore, there is a larger body of evidence around the

higher frequency brain waves like alpha, beta, and gamma.  

Gamma in particular has been studied rather thoroughly. Gamma waves are the most

common and appear in both sleeping and waking states (mech). They have been described in

multiple regions of the brain, but they are best described in the cortical structures where they are

best detected by EEGs (Gogia). These waves are the highest observed brain wave frequency and

are generally classified as ranging from about 30-100Hz. Interestingly, neurons can fire at a rate

as high as 1000 Hz (AI Impacts). So, it begs the question as to why the observed frequencies are

what they are. This suggests that inhibition is likely involved in regulation of brain wave

frequency.  

Considering that synchronization of neurons is more effective by IPSPs than EPSPs, two

primary models for gamma oscillations have been established: the I-I model and the E-I model

(Buzsáki). In both models, it is suspected that GABAA receptors establish the time constant to

create the gamma frequency because in recordings of membrane potentials of neurons firing at

gamma frequency the IPSPs cause the membrane to depolarize to able -60 mV which is the

equilibrium potential for Cl- (Buzsáki). In the I-I model it is inhibition of inhibitory interneurons

that allows EPSPs to fire in neuron pools, while in the E-I model there is only one level of

inhibition. In both cases, when GABAA receptor-mediated inhibition decays, the inhibited

neurons are able to spike again (Buzsáki). These models have been used to explain gamma

oscillations in the cortex but could be applied to other areas of the brain as well.  

While it appears to be well established that inhibition is mediated by GABAA receptors,

there are a few different receptors involved in the excitation of these neurons including

metabotropic glutamate receptors, muscarinic-cholinergic receptors, or kainate receptors

(Buzsáki). The cells that house these receptors include multipolar-pyramidal cells which act as

the primary excitation units while fast spiking PV basket cells are the primary inhibitory units

(Buzsáki).  There has been a variety of evidence pointing to PV basket cells as the inhibitory

units. For example, there is a correlation between spikes in these neurons and local field

potentials at gamma frequency in the hippocampus of rats (Buzsáki). Decreased gamma waves

also correlate with improperly functioning PV basket cells in schizophrenic individuals (Cell

Press). Another study found that rats that were genetically engineered to have deficient PV

basket cells had cognitive deficits that decreased performance on learning association tasks. In

the experiment rats were taught to associate food with a particular smell. Then the food was

moved to a container with a different smell. The genetically engineered rats were not as good at

learning the new association as the healthy controls. However, in a second experiment the

genetically engineered rats were treated with artificial stimulation to the prefrontal cortex at

gamma frequency, and the rats then performed as well as the healthy rats in the task up to a week

after receiving the stimulation. This suggests that PV cells are involved in creating gamma waves

and that gamma waves play a role in cognition and learning (Cell press).  

Another interesting aspect of brain waves is that frequencies in one area of the brain can

be coupled with the same or different frequencies in other areas of the brain.  This phenomenon

can occur in several ways. One frequency can be coupled with a different frequency, or an aspect

of one frequency in one region can determine an aspect of the wave frequency in another region.

For example, the phase of a frequency can determine the amplitude of a frequency in a different

region of the brain (Buzsáki).  

One type of frequency coupling is phase-phase coupling of gamma frequencies in the left

and right cerebral hemispheres. Coherence between gamma frequencies have been observed in
the left and right visual cortex as well as in homologous regions of the left and right

hippocampus and are mediated by interhemispheric axon tracts (Buzsáki). The distance between

these areas in the left and right hemispheres poses a problem for maintaining the same phase due

to conduction delays (Buzsáki). This problem can be alleviated by reciprocal connections

between the two regions which act as a feedback look to establish phase synchrony (Buzsáki). In

this case, the conduction delays still must be shorter than about 4-8 ms, and there need to

multiple cycles to allow synchrony to be established (Buzsáki).  

Brain waves can be either sinusoidal or nonsinusoidal. Nonsinusoidal brain waves have

typically been seen as a nuisance, and all spectral analysis methods used to calculate the

amplitude and phase of brain waves assume that brain waves can be characterized using a

sinusoidal basis (Cole). However, some neuroscientists argue that the presence of nonsinusoidal

wave form is significant and can be explained by underlying physiological mechanisms.  

To better understand these nonsinusoidal forms new methods for quantifying their

characteristics have had to be developed. One of the recently utilized methods is to quantify the

sharpness around peaks and troughs by determining the rate of change of voltage. The sharpness

measurements around peaks and troughs are then used to calculate a ratio between peak and

trough sharpness which acts as a measurement of peak and trough symmetry (Cole).  

A number of factors could be responsible for the shape of nonsinusoidal waveform.

However, few studies have been done that explain the physiological bases of these nonsinusoidal

waveforms. A few studies have suggested that waveform may be dependent on the

neurotransmitters present. For example, urethane anesthesia makes theta oscillations more

symmetric in the hippocampus, while kainate has been found to create sawtooth-shaped gamma
waves in the hippocampus (waveform). These studies suggest an underlying physiological

mechanism however more studies need to be done to come to conclusions.  

Though much has been learned about brain waves, there is still a lot of mystery behind

them. In particular the cellular mechanisms that create gamma oscillations in subcortical

structure are not well understood. However, the models created to describe them could serve as a

steppingstone to understand other possible mechanisms for creating gamma oscillations as well

as other brain wave frequencies. Understanding these mechanisms seems to be necessary for a

better understanding of frequency coupling and nonsinusoidal brain waves as well.  

 
 
 
 

 
 
 
 
 
 
 
 
References 
Buzsáki, György, and Xiao-Jing Wang. “Mechanisms of Gamma Oscillations.” Annual Review
of Neuroscience, vol. 35, no. 1, 2012, pp. 203–225., https://doi.org/10.1146/annurev-
neuro-062111-150444.  
Cole, Scott R., and Bradley Voytek. “Brain Oscillations and the Importance of Waveform
Shape.” Trends in Cognitive Sciences, vol. 21, no. 2, 2017, pp. 137–149.,
https://doi.org/10.1016/j.tics.2016.12.008.  
Gamma Rhythms, Interneurons, and Cognition, Cell Press, 18 Mar. 2015,
https://www.youtube.com/watch?app=desktop&v=dfNlodsd9Rw. Accessed 27 Nov.
2021.  
Lee, Darrin J., et al. “Review of the Neural Oscillations Underlying Meditation.” Frontiers in
Neuroscience, vol. 12, 2018, https://doi.org/10.3389/fnins.2018.00178.  
“Superhumans: The Remarkable Brain Waves of High-Level Meditators.” Daniel Goleman, Big
Think, https://bigthink.com/videos/daniel-goleman-superhumans-the-remarkable-brain-
waves-of-high-level-meditators/ 
Gogia AS, Martin Del Campo-Vera R, Chen KH, Sebastian R, Nune G, Kramer DR, Lee MB,
Tafreshi AR, Barbaro MF, Liu CY, Kellis S, Lee B. Gamma-band modulation in the
human amygdala during reaching movements. Neurosurg Focus. 2020 Jul;49(1):E4. doi:
10.3171/2020.4.FOCUS20179. PMID: 32610288. 
“Neuron Firing Rates in Humans.” AI Impacts, 11 Dec. 2020, https://aiimpacts.org/rate-of-
neuron-firing/.