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Seven papers on blink-related oscillations

“Reading notes from my AP Research annotated bibliography: what each paper did, what it found, and why it matters to the paper I'm writing.”

These are the seven sources from the annotated bibliography I turned in for AP Research on August 30, 2026, with the annotations as I wrote them. The topic is blink-related oscillations — the small burst of brain activity that follows every spontaneous blink — and what it might tell us about attention and mental workload.

First page of the submitted document
As submitted — August 30, 2026

01 — Blink-Related Delta Oscillations in the Resting-State EEG: A Wavelet Analysis.

Bonfiglio, Luca, et al. “Blink-Related Delta Oscillations in the Resting-State EEG: A Wavelet Analysis.” Neuroscience Letters, vol. 449, no. 1, 2009, pp. 57–60. doi.org/10.1016/j.neulet.2008.10.039

This article focuses on the electrical activity found in the brain after blinks. The authors were the first to detect delta-band waves after each blink. The authors used EEG machines to test this in seven participants who were at rest. Additionally, the authors provide explanations for the specific types of waves. The waves found looked similar to P300 waves, which tend to occur about 300 milliseconds after a trigger. However, these triggers are usually caused by external stimuli (exogenous). In this experiment, the P300 waves were caused by an internal stimulus (endogenous) of blinking which also is a natural and spontaneous action. Using the EEG, the authors were able to identify that the signals were in the medial parietal lobe. This was corroborated by Liu et al. when they found that the waves were specifically in the precuneus 8 years later. This article is of interest to me because it was one of the first studies to identify the delta blink-related oscillations.

02 — Cortical Source of Blink-Related Delta Oscillations and Their Correlation with Levels of Consciousness.

Bonfiglio, Luca, et al. “Cortical Source of Blink-Related Delta Oscillations and Their Correlation with Levels of Consciousness.” Human Brain Mapping, vol. 34, no. 9, 2013, pp. 2178–89. doi.org/10.1002/hbm.22056

This article focuses on the location of delta BROs in the cerebral cortex, and the correlation with levels of consciousness. The authors hypothesized that the delta BROs originated in the precuneus because they have been linked with continuous gathering of information from the surrounding environment. As stated in the paper, “From the functional point of view, the precuneus is involved in self-processing events with a visuospatial (attentional) connotation [Cavanna and Trimble, 2006], such as the continuous gathering of information from the surrounding environment and the representation of the self in relationship with the outside world [Gusnard and Raichle, 2001; Raichle et al., 2001]”. Therefore, they also hypothesized that the delta BROs would be absent or reduced in patients with disorders of consciousness (DOC). This is significant because fMRI studies have shown that precuneal activity is reduced or missing when consciousness is low or absent. In the experiment, the researchers recorded EEG activity at rest in both twelve healthy controls as well as nine patients with DOC (four vegetative states, and five minimally conscious states). By using Low Resolution Electromagnetic Tomography, the authors were able to estimate the location of the BROs in the precuneus. While the BROs were found in the precuneus in the control participants, delta BROs were not recognizable and no precuneal localization was possible in the patients with DOC. This will be useful in my paper, as it gives information on early studies on the location of delta BROs as well as on the correlation with cognitive function, as well as reinforcing that delta BROs reflect neural processes linked with awareness of the self and of the environment.

03 — The Origin and Purposes of Blinking.

Hall, Arthur. “The Origin and Purposes of Blinking.” The British Journal of Ophthalmology, vol. 29, no. 9, 1945, pp. 445–67.

This article focuses on the idea that blinking is not completely endogenous and can be affected by other factors. Hall believes that spontaneous blinking is also a way in which the brain stops visual fixation so that it can focus on readjusting to its surroundings. To begin, the author compared the blinking patterns of a control group to those of individuals with chronic encephalitis, which is a condition that affects high processing, and found that while individuals with encephalitis blink significantly less frequently during conversation (roughly 10.6 times per minute compared to 25.4), they blinked about the same amount as the control group while reading (roughly 2.9 times per minute compared to 3.3). Also of note was that both groups blinked much less when reading than during conversation, showing how there are differences in purposes of blinking. This was also shown in blinking patterns, as participants tended to blink at both physical gaps such as page turns, line changes, and column changes and at punctuation, rather than falling at random. Furthermore, blinks tend to occur at the exact moment when gaze shift from one object to another. Finally, even congenitally blind children blink at normal rates and even show reflex blinks in response to sudden sounds. While this source is over 80 years old, it is still useful for my research as it was the earliest demonstration of blink behavior's connections to neurological function and I will use it as a source that shows that spontaneous blinks tend to occur when attentional demand is low.

04 — Cognitive Loading via Mental Arithmetic Modulates Effects of Blink-Related Oscillations on Precuneus and Ventral Attention Network Regions.

Liu, Careesa C., et al. “Cognitive Loading via Mental Arithmetic Modulates Effects of Blink-Related Oscillations on Precuneus and Ventral Attention Network Regions.” Human Brain Mapping, vol. 40, no. 2, 2019, pp. 377–93. doi.org/10.1002/hbm.24378

This article focuses on the effects of cognitive loading on the brain activity linked with blink-related oscillations. These authors used magnetoencephalography to measure brain activity and brain wave location while participants were under cognitive load. They did this with two groups: one group performed mental arithmetic by counting down from 1000 by 7s, and the other group engaged in passive fixation. In both groups, the participants focused on an unchanging crosshair without any further sensory input. The study found that cognitive loading with mental arithmetic modulated BRO effects by decreasing cortical activations in the precuneus. However, in the mental arithmetic condition but not in the passive fixation, blinking activated regions of the ventral attention network (i.e., right supramarginal gyrus and inferior frontal gyrus), suggesting possible recruitment of these areas for blink processing under cognitive loading condition. The authors provide explanations for the effects of cognitive loading on cortical effects by suggesting a potential neurocognitive mechanism for blink processing in the precuneus. This will be useful in my paper as it provides important information for the effects of a stressor on the cognitive effects of BRO delta waves.

05 — Differential Neural Processing of Spontaneous Blinking under Visual and Auditory Sensory Environments: An EEG Investigation of Blink-Related Oscillations.

Liu, Careesa C., et al. “Differential Neural Processing of Spontaneous Blinking under Visual and Auditory Sensory Environments: An EEG Investigation of Blink-Related Oscillations.” NeuroImage, vol. 218, 2020, article 116879. doi.org/10.1016/j.neuroimage.2020.116879

This article focuses on the effects of visual and auditory sensory environments on BROs. The authors placed participants in environments with either no sensory stimulation, visual stimulation, or auditory stimulation. The blink-related oscillations were relatively unchanged in both the auditory and rest conditions, but changed under the visual condition. In the visual environment, there were two major differences. To begin, there were pre-blink spectral changes, which showed brain activity in anticipation of the blink. The authors relate this to memory, as these spectral theta waves were linked to more successful recall. In the visual condition, they believe that this brain activity took a mental snapshot before blinking to compare with the environment after the blink. Additionally, visual stimulation significantly reduced the BRO's amplitude. The researchers hypothesize that this was due to fewer neural resources left to process the blink-related visual information when the participants were already processing visual input. This information will be useful in my paper because it highlights the link between blinking and cognition, and further demonstrates the importance of BROs as a new window into brain function.

06 — Spontaneous Blinks Activate the Precuneus: Characterizing Blink-Related Oscillations Using Magnetoencephalography.

Liu, Careesa C., et al. “Spontaneous Blinks Activate the Precuneus: Characterizing Blink-Related Oscillations Using Magnetoencephalography.” Frontiers in Human Neuroscience, vol. 11, 2017, article 489. doi.org/10.3389/fnhum.2017.00489

This article focuses on the location of the small neural activity that comes every time that we blink while at rest. The authors believe that blinking triggers the precuneus in the brain. The precuneus is a central hub in the part of the brain that becomes active when you are not focused on the outside world, called the DMN, or Default Mode Network, and it is involved in visuo-spatial processing and awareness. While brain activity from blinking, often called BROs (blink-related oscillations) is usually tracked using electroencephalography, EEGs are not able to pinpoint local sources as well as MEGs. This study was the first to use MEG to confirm the location of the BROs. The authors used this magnetoencephalography to provide evidence for increased delta-band activity that peaks about 250 milliseconds after a blink which localizes to the bilateral precuneus. In the control condition, they tracked general magnetic activity in participants and in the blink condition they tracked the magnetic activity on the exact moment of a blink. This is of interest to me in my research because it is the first to confirm that blinks activate the precuneus using MEGs. This will provide background for how blink activity relates to brain function.

07 — Blink-Related Oscillations Provide Naturalistic Assessments of Brain Function and Cognitive Workload within Complex Real-World Multitasking Environments.

Page, Cleo, et al. “Blink-Related Oscillations Provide Naturalistic Assessments of Brain Function and Cognitive Workload within Complex Real-World Multitasking Environments.” Sensors, vol. 24, no. 4, 2024, article 1082. doi.org/10.3390/s24041082

This article focuses on whether blink-related oscillations can be used as predictors of cognitive function in pilots. The authors believe that because BROs are spontaneous endogenous behaviors, they could be used to measure a person's mental workload without interrupting them with outside stimuli. To test this, the researchers looked at EEG data from eight adult participants that completed the Multi-Attribute Task Battery (MATB), a NASA task that simulates benchmarked complex and realistic tasks analogous to those undertaken by pilots/aircrews in flight, at low, medium, and high workloads. The study found that BROs were present at every workload level, demonstrating that BROs appear even while multitasking. The BRO was correlated with mental workload in several ways, such as fewer blinks at more difficulty, decreased amplitude in the back of the head, increased amplitude in the parietal lobe, and a pre-blink theta change in blinks under high workload. The authors used this to show that BROs can passively capture cognitive loading effects. This will be useful in my paper as one of the possible future uses for tracking blink-related oscillations.