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Unihemispheric slow-wave sleep

By Kimberly Mauro

Introduction

Unihemispheric slow-wave sleep is an extraordinary phenomenon in which half of the brain is

asleep while the other half is awake. The term “unihemispheric” comes from the Latin prefix

uni- meaning “one” and hemisphere meaning “half”. In unihemispheric slow-wave sleep,

abbreviated USWS, half of the brain is sleeping with the opposite side eye open. The other eye is

closed, corresponding to the other side of the brain that is awake. In other words, the eye-to-brain

correlation is crossed like an “X”, called an optic chiasm. This unique form of sleep is different

from sleep in humans and most land animals in which both halves of the brain are resting with

closed eyes. USWS has been observed in aquatic mammals such as dolphins and whales, the

Northern fur seal, and various species of birds and is the ultimate life-saver in times of predation.

REM & NREM Sleep

USWS is a form of NREM (non-rapid eye movement) sleep that exhibits slow brain waves and is

commonly known as deep sleep. Both humans and animals have NREM sleep, but in USWS,

NREM only occurs in the resting hemisphere. NREM sleep uses very little energy and is

important in repairing muscle, bone, and tissue, restoring energy, and strengthening metabolic
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processes in the body (Matheny, 2020). In contrast, REM (rapid eye movement) sleep involves

high brain activity in which we experience dreams. Birds exhibit USWS during specific times

discussed later in this paper, but can also undergo REM sleep. Marine mammals like dolphins

and whales only experience USWS and do not experience any other kind of sleep. The Northern

fur seal can surprisingly alternate between USWS in the water and REM sleep only while on

land in which they sleep with both halves of the brain (Kendall-Bar et al., 2019). Switching sleep

types based on location has not been seen in other USWS animals.

Mechanism

Production of Acetylcholine

Acetylcholine is a chemical in the brain that promotes wakefulness and activates the hemispheres

in the brain. During REM sleep and normal sleep conditions, acetylcholine is released in the

same amounts in each hemisphere. However, research shows that during USWS, the hemisphere

that is in deep sleep will have significantly less acetylcholine released than the side that is awake.

This process is called lateralized acetylcholine release (Lapierre, 2007). Lateralization is the

tendency for neurons in the brain to function on one side of the brain in a specialized fashion.

Lateralization in acetylcholine production could indicate that each hemisphere has specialized

neurons to carry out specific functions and keep half of the brain awake while the other can be

temporarily “shut down”.

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Temperature Effects

Research shows that brain temperatures in dolphins and whales decrease while in USWS.

Although there is no distinct cause for temperature changes, this reaction could be correlated

with thermoregulation. Thermoregulation can enable muscular movement which may prevent

their bodies from getting too cold and enables breathing since dolphins and whales need to be

awake to breathe in air periodically (Lyamin, 2017).

Hemisphere to Eye Correlation

One commonly observed feature in animals exhibiting USWS is

the correlation between eye and hemisphere. If you have ever

heard the phrase, “sleep with one eye open”, that is exactly what

these animals do. When in USWS, one eye is open while the

other eye remains closed. The eye that is open is found to be

connected to the hemisphere that is awake on the opposite side of

the body and the closed eye connects to the sleeping hemisphere

on the other side in a cross-like pattern. Eye-opening plays an

amazing role in keeping the hemisphere awake, but is considered
to be a side effect of USWS and not the major player since
Figure 1. Greater Racket-Tailed Drongo in
Unihemispheric Slow-wave Sleep by BESG, 2015,
https://besgroup.org/2015/07/07/greater-racket-tailed-
drongo-in-unihemispheric-slow-wave-sleep/

USWS can occur in blind animals such as the Indus river dolphin

and blind chickens (Mascetti, 2016). This puzzling phenomenon

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conflicts with the need for animals with predation pressures to

perform USWS and has no recent solutions to why blind animals

may have USWS. Studies propose that eye to hemisphere

correlation can be preferential in cases of threatening

environments or to conserve energy.

Anatomical Differences

Corpus Callosum

Humans, land mammals, and animals that experience USWS all have a corpus callosum, a nerve

bundle located in the middle of the brain and is responsible for communication between the two

hemispheres. Marine mammals have a smaller corpus callosum compared to humans and other

mammals and birds entirely lack a corpus callosum. Since it is necessary for USWS to have each

hemisphere functioning somewhat independently of each other, scientists predict that a smaller

corpus callosum could result in less communication between the halves of the brain (Beaumont,

2019). This speculation could be why the halves of the brain can function independently in

USWS. If there is reduced communication between the hemispheres, then this might explain how

USWS animals are able to sleep and be awake at the same time, but scientists have yet to

determine if a smaller corpus callosum relates to brain functioning in USWS.

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Locus Coeruleus

The locus coeruleus is a portion of the brainstem that synthesizes norepinephrine, a hormone in

the brain that increases arousal during times of stress in the fight-or-flight response. During

USWS, norepinephrine is asymmetrically produced with high levels in the conscious hemisphere

and low levels in the resting hemisphere. Similarly, humans release low levels of norepinephrine

while sleeping and increasing levels while waking up (“Norepinephrine”, 2020). Based on

anatomical observations, mammals that demonstrate USWS have a larger posterior commissure

containing fibers that connect both hemispheres. The fibers from the locus coeruleus in animals

that have USWS are more crossed and entangled in the brainstem than in individuals that do not

experience USWS which could help regulate the sleep-wake cycles in USWS.

Tracing USWS in the Brain

EEG Usage
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Imaging techniques in modern medicine allow scientists to detect and record electrical activity

in the brain using an EEG test. An EEG is an advanced piece of technology composed of a

series of electrical discs and wires that attach to the surface of the head and measure electrical

currents in the brain with no surgical means. Doctors commonly perform this impressive test

since it does not require an internal operation or recovery time. Electrical currents are flowing

through the brain all the time whether you are awake or asleep; this is the way our brain cells

communicate (Mayo Clinic, 2020). Since fur seals can switch from normal REM sleep also

called bilateral slow-wave sleep (BSWS) and USWS, EEGs provide a great presentation of the

different brain waves in each hemisphere. In BSWS, the brain

waves between the left and right hemisphere are the same,

while in USWS, brain waves are more prominent on the left or

right hemisphere depending on the animal and the environment

it is in (Figure 2).

Figure 2. An EEG test on the Northern fur seal

Behaviors exhibiting BSWS and USWS, also called ASWS
in the left and right hemisphere by Lapierre et al.,
2007, Journal of Neuroscience,
https://www.jneurosci.org/content/27/44/11999

Group Edge Effect in Birds

When birds are in a group called a flock, typically they fly in a V formation or sleep close to

one another with some birds in the center and others along the edge in a process called the

“group edge effect”. The Mallard duck is a prime example of an animal that while in this
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formation, performs USWS. The ducks on the outside edge of the group will sleep under USWS

with the open eye pointing away from the group and towards the outside edge. The center ducks

usually undergo BSWS and do not have USWS while at the center. In this formation, the ducks

can watch out for predators while half asleep. In flight, birds along the edge experience USWS

to stay alert of their surroundings and are also thought to conserve energy during long migration

periods by resting half of their brain. EEG tests have confirmed USWS in the “group edge

effect” (Rattenborg, 1999).

Fur Seal Sleep Posture

When in the water, Northern fur seals reveal a lateral posture on the surface during USWS. In this

position, their back flippers stay above the water and one front flipper is outside of the water. The other

front flipper is submerged below the surface and paddles to stabilize their body in this position. The

open eye is in the water correlating to the active hemisphere and moving flipper, while the closed eye

is above the surface (Kendall-Bar et al., 2019). This remarkable behavior is not seen in many aquatic

mammals which suggest that sleep behaviors are

specific to each species and can be deemed

preferential. A large assumption is that because

Northern fur seals are under more pressure from

predators such as killer whales and great white

sharks, there is a larger demand for USWS to avoid

Figure 3. Sleep behavior of a northern fur seal at sea by Ungurean, G.,
such predators (Lyamin et al., 2017). & Rattenborg, N. C., 2018, https://www.cell.com/current-
biology/fulltext/S0960-9822(18)30608-0
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References

Beaumont, P. (2019, October 5). Sleep, but not as we know it. Retrieved from

https://www.seawatchfoundation.org.uk/sleep-but-not-as-we-know-it/

BESG. (2015, July 07). Greater Racket-tailed Drongo in Unihemispheric Slow-wave Sleep.

Retrieved from https://besgroup.org/2015/07/07/greater-racket-tailed-drongo-in-

unihemispheric-slow-wave-sleep/

EEG (electroencephalogram). (2020, April 15). Retrieved from

https://www.mayoclinic.org/tests-procedures/eeg/about/pac-20393875

Kendall-Bar, J. M., Vyssotski, A. L., Mukhametov, L. M., Siegel, J. M., & Lyamin, O. I. (2019).

Eye state asymmetry during aquatic unihemispheric slow wave sleep in northern fur seals

(Callorhinus ursinus). Plos One, 14(5). doi:10.1371/journal.pone.0217025

Lapierre, J., Kosenko, P., Lyamin, O., Kodama, T., Mukhametov, L., & Siegel, J. (2007, October

31). Cortical Acetylcholine Release Is Lateralized during Asymmetrical Slow-Wave Sleep

in Northern Fur Seals. Retrieved from https://www.jneurosci.org/content/27/44/11999

Lateralization of brain function. (2020, September 17). Retrieved from

https://en.wikipedia.org/wiki/Lateralization_of_brain_function

Lyamin, O., Mukhametov, L., & Siegel, J. (2017, June). Sleep in the northern fur seal. Retrieved

from https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5609733/
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Mascetti, G. (2016, July 12). Unihemispheric sleep and asymmetrical sleep: Behavioral,

neurophysiological, and functional perspectives. Retrieved from

https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4948738/

Matheny, S. (2020, March 26). Why Do We Sleep? - The Importance of REM & NREM.

Retrieved from https://www.nordicnaturals.com/healthy-science/what-happens-when-we-

sleep/

Norepinephrine. (2020, September 17). Retrieved from

https://en.wikipedia.org/wiki/Norepinephrine

Rattenborg, N. C., Lima, S. L., & Amlaner, C. J. (1999, February). Half-awake to the risk of

predation. Nature, 397(6718), 397-398. doi:10.1038/17037

Ungurean, G., & Rattenborg, N. C. (2018, June 18). Neuroethology: Fur Seals Don't Lose Sleep

over REM Lost at Sea. Retrieved from https://www.cell.com/current-

biology/fulltext/S0960-9822(18)30608-0

Unihemispheric slow-wave sleep. (2020, August 17). Retrieved from

https://en.wikipedia.org/wiki/Unihemispheric_slow-wave_sleep

In this paper, I utilized Markel’s strategies of comparison and contrast, graphics,

partitioning, examples, and etymology to define my term in greater effect. Comparison

and contrast were used in the “Anatomical Differences” subdivision on page 3 to

compare the anatomy of humans and animals that sleep in BSWS with animals that sleep
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in USWS. Graphics are also an important aspect of my paper found on pages 3, 4, and 5

to elaborate on the paragraph adjacent to it in a visual. Partitioning was implemented in

page 2 under the “Mechanism” subdivision to break down portions of how USWS works

in the brain. Examples are found on page 5 under the “Behavior” subdivision to

exemplify certain behaviors in different species. Etymology was one of the first strategies

given on page 1 under the first subdivision, “Introduction” to reveal how the definition

derived to enhance the meaning of the definition.