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Circadian rhythm
From Wikipedia, the free encyclopedia

A circadian rhythm is an approximate daily periodicity, a roughly-24-hour cycle in the biochemical,

physiological or behavioural processes of living beings, including plants, animals, fungi and
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cyanobacteria. The term "circadian", coined by Franz Halberg,[1] comes from the Latin circa, "around",
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and diem or dies, "day", meaning literally "approximately one day." The formal study of biological
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temporal rhythms such as daily, tidal, weekly, seasonal, and annual rhythms, is called chronobiology.
Featured content
Current events Circadian rhythms are endogenously generated, and can be entrained by external cues, called
Random article Zeitgebers. The primary one is daylight. These rhythms allow organisms to anticipate and prepare for
precise and regular environmental changes.
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Contents [hide]
Go Search
1 History
2 Criteria
interaction
3 Origin
About Wikipedia
4 Importance in animals
Community portal
4.1 Impact of light-dark cycle
Recent changes
4.2 Arctic animals
Contact Wikipedia
5 The biological clock in mammals
Donate to Wikipedia
5.1 Determining the human circadian rhythm
Help
5.2 Outside the "master clock"
toolbox 6 Light and the biological clock
What links here 6.1 The myth of the 25-hour day
Related changes 6.2 The human circadian period
Upload file 7 Human health
Special pages 7.1 Disruption
Printable version 8 Effects on cocaine sensitization in mice
Permanent link
9 See also
Cite this page
10 Notes
languages 11 Further reading
Deutsch 12 External links
Español
Français
Italiano
‫עברית‬ History [edit]

Nederlands
The first endogenous circadian oscillation was observed in the 1700s by the French scientist Jean-
日本語
Jacques d'Ortous de Mairan who noticed that 24-hour patterns in the movement of the leaves of the
Norsk (nynorsk)
plant Mimosa pudica continued even when the plants were isolated from external stimuli.
Polski
Português In 1918 J. S. Szymanski showed that animals are capable of maintaining 24-hour activity patterns in the
Русский absence of external cues such as light and changes in temperature.[2]
Српски / Srpski
The earliest known account of a circadian rhythm dates from the 4th century BC, when Androsthenes,
Svenska
in descriptions of the marches of Alexander the Great, described diurnal leaf movements of the
tamarind tree.

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Criteria [edit]

Three general criteria of circadian rhythms are necessary to differentiate genuinely endogenous rhythms
from coincidental or apparent ones: the rhythms persist in the absence of cues, they can be brought to
match the local time, and will do so in a precise manner over a range of temperatures.
The rhythm persists in constant conditions (for example, constant dark) with a period of about 24
hours. The rationale for this criterion is to distinguish circadian rhythms from those "apparent"
rhythms which merely are responses to external periodic cues. A rhythm cannot be declared to be
endogenous unless it has been tested in conditions without external periodic input.
The rhythm is temperature-compensated, i.e. it maintains the same period over a range of
temperatures. The rationale for this criterion is to distinguish circadian rhythms from other biological
rhythms arising due to the circular nature of a reaction pathway. At a low enough or high enough
temperature, the period of a circular reaction may reach 24 hours, but it will be merely coincidental.
The rhythm can be reset by exposure to an external stimulus. The rationale for this criterion is to
distinguish circadian rhythms from other imaginable endogenous 24-hour rhythms that are immune
to resetting by external cues and hence do not serve the purpose of estimating the local time. Travel
across time zones illustrates the necessity of the ability to adjust the biological clock so that it can
reflect the local time and anticipate what will happen next.

Origin [edit]

This section does not cite any references or sources.

Please help improve this section by adding citations to reliable sources.
Unverifiable material may be challenged and removed. (October 2007)

Photosensitive proteins and circadian rhythms are believed to have originated in the earliest cells, with
the purpose of protecting the replicating of DNA from high ultraviolet radiation during the daytime. As a
result, replication was relegated to the dark. The fungus Neurospora, which exists today, retains this
clock-regulated mechanism. Rhythmicity appears to be as important in regulating cyclic biochemical
processes within an individual, as in coordinating with the environment. This is suggested by the
maintenance (heritability) of circadian rhythms in fruit flies after several hundred generations in constant
laboratory conditions (Sheeba et al. 1999), as well as the experimental elimination of behavioral but not
physiological circadian rhythms in quail (Guyomarc'h et al. 1998, Zivkovic et al. 1999).
The simplest known circadian clock is that of the prokaryotic cyanobacteria. Recent research has
demonstrated that the circadian clock of Synechococcus elongatus can be reconstituted in vitro with
just the three proteins of their central oscillator. This clock has been shown to sustain a 22-hour rhythm
over several days upon the addition of ATP. Previous explanations of the prokaryotic circadian
timekeeper were dependent upon a DNA transcription / translation feedback mechanism. It is an
outstanding question whether circadian clocks in eukaryotic organisms require translation/transcription-
derived oscillations. For although the circadian systems of eukaryotes and prokaryotes have the same
basic architecture: input - central oscillator - output, they do not share any homology. This implies
probable independent origins.
In 1971, Ronald J. Konopka and Seymour Benzer first identified a genetic component of the biological
clock using the fruit fly as a model system. Three mutant lines of flies displayed aberrant behaviour -
one had a shorter period, another had a longer one and the third had none. All three mutations mapped
to the same gene, which was named period. [3] The same gene was identified to be defective in the
sleep disorder FASPS (Familial Advanced Sleep Phase Syndrome) in human beings thirty years later -
underscoring the conserved nature of the molecular circadian clock through evolution. We now know
many more genetic components of the biological clock. Their interactions result in an interlocked

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feedback loop of gene products resulting in periodic fluctuations that the cells of the body interpret as a
specific time of the day.
A great deal of research on biological clocks was done in the latter half of the 20th century. It is now
known that the molecular circadian clock can function within a single cell; i.e., it is cell-autonomous. [4]
At the same time, different cells may communicate with each other resulting in a synchronised output
of electrical signaling. These may interface with endocrine glands of the brain to result in periodic
release of hormones. The receptors for these hormones may be located far across the body and
synchronise the peripheral clocks of various organs. Thus, the information of the time of the day as
relayed by the eyes travels to the clock in the brain, and, through that, clocks in the rest of the body
may be synchronised. This is how the timing of, for example, sleep/wake, body temperature, thirst, and
appetite are coordinately controlled by the biological clock.

Importance in animals [edit]

Circadian rhythms are important in determining the sleeping and feeding patterns of all animals,
including human beings. There are clear patterns of core body temperature, brain wave activity,
hormone production, cell regeneration and other biological activities linked to this daily cycle. In
addition, photoperiodism, the physiological reaction of organisms to the length of day or night, is vital to
both plants and animals, and the circadian system plays a role in the measurement and interpretation
of daylength.

«Timely prediction of seasonal periods of weather conditions, food availability or predator activity is crucial
for survival of many species. Although not the only parameter, the changing length of the photoperiod
('daylength') is the most predictive environmental cue for the seasonal timing of physiology and behavior,
most notably for timing of migration, hibernation and reproduction.»[5]

Impact of light-dark cycle [edit]

The rhythm is linked to the light-dark cycle. Animals, including humans, kept in total darkness for
extended periods eventually function with a freerunning rhythm. Each "day," their sleep cycle is pushed
back or forward, depending on whether their endogenous period is shorter or longer than 24 hours. The
environmental cues that each day reset the rhythms are called Zeitgebers (from the German, Time
Givers).[6] It is interesting to note that totally-blind subterranean mammals (e.g., blind mole rat Spalax
sp.) are able to maintain their endogenous clocks in the apparent absence of external stimuli.
Freerunning organisms that normally have one consolidated sleep episode will still have it when in an
environment shielded from external cues, but the rhythm is, of course, not entrained to the 24-hour
light/dark cycle in nature. The sleep/wake rhythm may, in these circumstances, become out of phase
with other circadian or ultradian rhythms such as temperature and digestion.[citation needed]
Recent research has influenced the design of spacecraft environments, as systems that mimic the
light/dark cycle have been found to be highly beneficial to astronauts.

Arctic animals [edit]

Norwegian researchers at the University of Tromsø have shown that some Arctic animals (ptarmigan,
reindeer) show circadian rhythms only in the parts of the year that have daily sunrises and sunsets. In
one study of reindeer, animals at 70 degrees North showed circadian rhythms in the autumn, winter,
and spring, but not in the summer. Reindeer at 78 degrees North showed such rhythms only autumn
and spring. The researchers suspect that other Arctic animals as well may not show circadian rhythms
[7][8]

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in the constant light of summer and the constant dark of winter.

However, another study in northern Alaska found that ground squirrels and porcupines strictly
maintained their circadian rhythms through 82 days and nights of sunshine. The researchers speculate
that these two small mammals see that the apparent distance between the sun and the horizon is
shortest once a day, and, thus, a sufficient signal to adjust by. [9]

The biological clock in mammals [edit]

The primary circadian "clock" in mammals is located in the suprachiasmatic nucleus (or nuclei) (SCN),
a pair of distinct groups of cells located in the hypothalamus. Destruction of the SCN results in the
complete absence of a regular sleep/wake rhythm. The SCN receives information about illumination
through the eyes. The retina of the eyes contains not only "classical" photoreceptors but also
photoresponsive retinal ganglion cells. These cells, which contain a photo pigment called melanopsin,
follow a pathway called the retinohypothalamic tract, leading to the SCN. If cells from the SCN are
removed and cultured, they maintain their own rhythm in the absence of external cues.
It appears that the SCN takes the information on day length from the retina, interprets it, and passes it
on to the pineal gland, a tiny structure shaped like a pine cone and located on the epithalamus. In
response the pineal secretes the hormone melatonin. Secretion of melatonin peaks at night and ebbs
during the day.
The circadian rhythms of humans can be entrained to slightly shorter and longer periods than the
earth's 24 hours. Researchers at Harvard have recently shown that human subjects can at least be
entrained to a 23.5-hour cycle and a 24.65-hour cycle (the latter being the natural solar day-night cycle
on the planet Mars).[10]

Determining the human circadian rhythm [edit]

The classic phase markers for measuring the

timing of a mammal's circadian rhythm are
melatonin secretion by the pineal gland
and core body temperature.
For temperature studies, people must remain
awake but calm and semi-reclined in near
darkness while their rectal temperatures are
taken continuously. The average human adult's
temperature reaches its minimum at about
Overview of human circadian biological clock with some
05:00 (5 a.m.), about two hours before habitual
physiological parameters.
wake time, though variation is great among
normal chronotypes.
Melatonin is absent from the system or undetectably low during daytime. Its onset in dim light, dim-light
melatonin onset (DLMO), at about 21:00 (9 p.m.) can be measured in the blood or the saliva. Both
DLMO and the midpoint (in time) of the presence of the hormone in the blood or saliva have been used
as circadian markers.
However, newer research indicates that the melatonin offset may be the most reliable marker. Benloucif
et al in Chicago in 2005 found that melatonin phase markers were more stable and more highly
correlated with the timing of sleep than the core temperature minimum. They found that both sleep
offset and melatonin offset were more strongly correlated with the various phase markers than sleep
onset. In addition, the declining phase of the melatonin levels was more reliable and stable than the
termination of melatonin synthesis.[11]

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One method used for measuring melatonin offset is to analyze a sequence of urine samples throughout
the morning for the presence of the melatonin metabolite 6-sulphatoxymelatonin (aMT6s). Laberge et al
in Quebec in 1997 used this method in a study which confirmed the frequently found delayed circadian
phase in healthy adolescents.[12]

Outside the "master clock" [edit]

More-or-less independent circadian rhythms are found in many organs and cells in the body outside
the suprachiasmatic nuclei (SCN), the "master clock." These clocks, called peripheral oscillators, are
found in the esophagus, lung, liver, pancreas, spleen, thymus and the skin.[13] Though oscillators in the
skin respond to light, a systemic influence has not been proven so far.[14][15] There is some evidence
that also the olfactory bulb and prostate may experience oscillations when cultured, suggesting that also
these structures may be weak oscillators.
Furthermore, liver cells, for example, appear to respond to feeding rather than to light. Cells from many
parts of the body appear to have freerunning rhythms.

Light and the biological clock [edit]

Light resets the biological clock in accordance with the phase response curve (PRC). Depending on the
timing, light can advance or delay the circadian rhythm. Both the PRC and the required illuminance
vary from species to species; much lower light levels are required to reset the clocks in nocturnal
rodents than in humans.
In addition to light intensity, wavelength (or color) of light is an important factor in the degree to which
the clock is reset. Melanopsin is most efficiently excited by blue light, 420-440 nm [16] according to
some researchers while others have reported 470-485nm.

The myth of the 25-hour day [edit]

Early investigators determined the human circadian period to be 25 hours or more. They went to great
lengths to shield subjects from time cues and daylight, but they were not aware of the effects of indoor
electric lights. The subjects were allowed to turn on light when they were awake and to turn it off when
they wanted to sleep. Electric light in the evening delayed their circadian phase. These results became
well known. [17]

The human circadian period [edit]

Modern research under very controlled conditions has shown the human period for adults to be just
slightly longer than 24 hours on average. Czeisler et al at Harvard found the range for normal, healthy
adults of all ages to be quite narrow: 24 hours and 11 minutes ± 16 minutes. The "clock" resets itself
daily to the 24-hour cycle of the earth's rotation.[17]

Human health [edit]

Timing of medical treatment in coordination with the body clock may significantly increase efficacy and
reduce drug toxicity or adverse reactions. For example, appropriately timed treatment with angiotensin
converting enzyme inhibitors (ACEi) may reduce nocturnal blood pressure and also benefit left
ventricular (reverse) remodeling. There are many health problems associated with a disturbance in the

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human circadian rhythm, such as Seasonal Affective Disorder (SAD), delayed sleep phase syndrome
(DSPS) and other circadian rhythm disorders.[18] Circadian rhythms also play a part in the reticular
activating system which is crucial for maintaining a state of consciousness. In addition, a reversal in the
sleep-wake cycle may be a sign or complication of uremia,[19] azotemia or acute renal failure.

Disruption [edit]

Disruption to rhythms usually has a negative effect. Many travelers have experienced the condition
known as jet lag, with its associated symptoms of fatigue, disorientation and insomnia.
A number of other disorders, for example bipolar disorder and some sleep disorders, are associated
with irregular or pathological functioning of circadian rhythms. Recent research suggests that circadian
rhythm disturbances found in bipolar disorder are positively influenced by lithium's effect on clock
genes. [20]
Disruption to rhythms in the longer term is believed to have significant adverse health consequences on
peripheral organs outside the brain, particularly in the development or exacerbation of cardiovascular
disease. The suppression of melatonin production associated with the disruption of the circadian rhythm
may increase the risk of developing cancer.[21][22]

Effects on cocaine sensitization in mice [edit]

Circadian rhythms and clock genes expressed in brain regions outside the SCN may significantly
influence the effects produced by drugs such as cocaine. [23][24] Moreover, genetic manipulations of
clock genes profoundly affect cocaine's actions.[25]

See also [edit]

Actigraphy (also known as Actimetry)

Advanced sleep phase syndrome
ARNTL
ARNTL2
Chronobiology
Chronotype
Circadian rhythm sleep disorders
Cryptochrome
CRY1 and CRY2, the cryptochrome family genes
Jet lag
Light effects on circadian rhythm
PER1, PER2, and PER3, the period family genes

Notes [edit]

1. ^ http://www.msi.umn.edu/~halberg Halberg Chronobiology Centre

2. ^ Danchin, Antoine. "Important dates 1900-1919 ". HKU-Pasteur Research Centre. Retrieved on 2008-01-12.
3. ^ Purves, Dale et al [2001]. "Molecular Mechanisms of Biological Clocks ", NEUROSCIENCE (e-book), second
edition, Sunderland, MA, U.S.A.: Sinauer Associates. ISBN 0-87893-742-0. Retrieved on 2008-05-30.
4. ^ Circadian gene expression in individual fibroblast...[Cell. 2004] - PubMed Result
5. ^ Zivkovic, Bora "Coturnix" (2005-08-13 / July 25, 2007). "Clock Tutorial #16: Photoperiodism - Models and

http://en.wikipedia.org/wiki/Circadian_rhythm[30.09.2008 19:04:44]
Circadian rhythm - Wikipedia, the free encyclopedia

Experimental Approaches ". A Blog Around the Clock. ScienceBlogs. Retrieved on 2007-12-09.
6. ^ "Circadian rhythms ". Rapid eye movement (REM) sleep. Armenian Medical Network (2007). Retrieved on
2007-09-19.
7. ^ Spilde, Ingrid (December 2005). "Reinsdyr uten døgnrytme " (in Language: Norwegian, Bokmål), forskning.no.
Retrieved on 2007-11-24.
8. ^ Zivkovic, Bora, aka Coturnix, chronobiologist. "Circadian Rhythms, or Not, in Arctic Reindeer ". A Blog around
the Clock. ScienceBlogs.com. Retrieved on 2007-11-24.
9. ^ Zivkovic, Bora, aka Coturnix, chronobiologist (2007-02-11). "Small Arctic Mammals Entrain to Something during
the Long Summer Day ". A Blog Around the Clock. ScienceBlogs.com. Retrieved on 2007-11-26.
10. ^ Scheer, Frank A. J. L.; Kenneth P. Wright, Jr., Richard E. Kronauer, Charles A. Czeisler (2007-08-08). "Plasticity
of the Intrinsic Period of the Human Circadian Timing System ". PLoS ONE 2: e721.
doi:10.1371/journal.pone.0000721 . Retrieved on 2007-12-31.
11. ^ Benloucif, S.; Guico, M.J.; Reid, K.J.; Wolfe, L.F.; L'Hermite-Baleriaux, M.; Zee, P.C. (2005). "Stability of
melatonin and temperature as circadian phase markers and their relation to sleep times in humans ". J Biol
Rhythms 20 (2): pages 178–88. Chicago, Illinois, USA: Center for Sleep and Circadian Biology, Departments of
Neurology, Northwestern University Feinberg School of Medicine. doi:10.1177/0748730404273983 . PMID
15834114 . Retrieved on 2007-12-18.
12. ^ Laberge, L.; Lesperance, P.; Tremblay, R.; Lambert, C.; Montplaisir, J. (1997). "Phase delay of 6-
sulphatoxymelatonin in normal adolescents " (in English). Sleep Research 26: p. 727. Québec, Canada: Centre
d'etude du Sommeil, Hopital du Sacre-Coeur, Département de Psychologie, Département de Pharmacologie,
Departement de Psychiatrie, Université de Montréal. Retrieved on 2007-12-18.
13. ^ Zanello, S. et al., J. Inv. Dermatol. 2000, Vol. 115, 4 Oct.: Expression of the Circadian Clock Genes clock and
period1 in Human Skin
14. ^ Kawara, S. et al. J. Inv. Derm. 2002, Vol 119, 6 Dec.: Low-dose UVB Rays Alter the mRNA Expression of the
Circadian Clock Genes in cultured Human Keratinocytes
15. ^ Campbell, S. and Murphy, P, Science 1998, Vol 279, 16 Jan.: Extraocular Circadian Phototransduction in Humans
16. ^ Newman LA, Walker MT, Brown RL, Cronin TW, Robinson PR: "Melanopsin forms a functional short-wavelength
photopigment", Biochemistry. 2003 Nov 11;42(44):12734-8.
17. ^ a b "Human Biological Clock Set Back an Hour " (1999). Retrieved on 2007-09-23. "The variation between our
subjects, with a 95 percent level of confidence, was no more than plus or minus 16 minutes, a remarkably small
range."
18. ^ "Circadian Rhythms and Sleep ". Circadian Rhythms and Sleep. Serendip (2007). Retrieved on 2007-09-19.
19. ^ Sinert, Richard; Peter R Peacock, Jr (May 10, 2006). "Renal Failure, Acute ". eMedicine from WebMD.
Retrieved on 2008-08-03.
20. ^ NIMH · Science News from 2006 · Lithium Blocks Enzyme To Help Cells’ Clocks Keep On Tickin’
21. ^ Straif K, Baan R, Grosse Y, Secretan B, El Ghissassi F, Bouvard V, Altieri A, Benbrahim-Tallaa L, Cogliano V,
WHO International Agency for Research on Cancer Monograph Working Group. Carcinogenicity of shift-work,
painting, and fire-fighting. [1] Lancet Oncol. 2007; 12(8):1065-1066.
22. ^ WebMD: Night Shift Work May Cause Cancer
23. ^ Uz T, Akhisaroglu M, Ahmed R, Manev H (2003). "The pineal gland is critical for circadian Period1 expression in
the striatum and for circadian cocaine sensitization in mice". Neuropsychopharmacology 28 (12): 2117–23.
doi:10.1038/sj.npp.1300254 . PMID 12865893 .
24. ^ Kurtuncu M, Arslan A, Akhisaroglu M, Manev H, Uz T (2004). "Involvement of the pineal gland in diurnal cocaine
reward in mice". Eur J Pharmacol 489 (3): 203–5. doi:10.1016/j.ejphar.2004.03.010 . PMID 15087244 .
25. ^ McClung C, Sidiropoulou K, Vitaterna M, Takahashi J, White F, Cooper D, Nestler E (2005). "Regulation of
dopaminergic transmission and cocaine reward by the Clock gene". Proc Natl Acad Sci USA 102 (26): 9377–81.
PMID 15967985 .

Further reading [edit]

Aschoff J (ed.) (1965) Circadian Clocks. North Holland Press, Amsterdam

Avivi A, Albrecht U, Oster H, Joel A, Beiles A, Nevo E. 2001. Biological clock in total darkness: the
Clock/MOP3 circadian system of the blind subterranean mole rat. Proc Natl Acad Sci USA
98:13751- 13756.

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Circadian rhythm - Wikipedia, the free encyclopedia

Avivi A, Oster H, Joel A, Beiles A, Albrecht U, Nevo E. 2002. Circadian genes in a blind
subterranean mammal II: conservation and uniqueness of the three Period homologs in the blind
subterranean mole rat, Spalax ehrenbergi superspecies. Proc Natl Acad Sci USA 99:11718-11723.
Ditty JL, Williams SB, Golden SS (2003) A cyanobacterial circadian timing mechanism. Annu Rev
Genet 37:513-543
Dunlap JC, Loros J, DeCoursey PJ (2003) Chronobiology: Biological Timekeeping. Sinauer,
Sunderland
Dvornyk V, Vinogradova ON, Nevo E (2003) Origin and evolution of circadian clock genes in
prokaryotes. Proc Natl Acad Sci USA 100:2495-2500
Koukkari WL, Sothern RB (2006) Introducing Biological Rhythms. Springer, New York
Martino T, Arab S, Straume M, Belsham DD, Tata N, Cai F, Liu P, Trivieri M, Ralph M, Sole MJ.
Day/night rhythms in gene expression of the normal murine heart. J Mol Med. 2004 Apr;82(4):256-
64. Epub 2004 Feb 24. PMID: 14985853
Refinetti R (2006) Circadian Physiology, 2nd ed. CRC Press, Boca Raton
Takahashi JS, Zatz M (1982) Regulation of circadian rhythmicity. Science 217:1104–1111
Tomita J, Nakajima M, Kondo T, Iwasaki H (2005) No transcription–translation feedback in circadian
rhythm of KaiC phosphorylation. Science 307: 251–254
Moore-Ede, Martin C., Sulszman, Frank M., and Fuller, Charles A. (1982) "The Clocks that Time Us:
Physiology of the Circadian Timing System." Harvard University Press, Cambridge, MA. ISBN 0-674-
13581-4.

External links [edit]

Can you reset your body clock? by John Triggs in the Daily Express December 4 2007
Circadian rhythm at the Open Directory Project
www.circadian.org
Biological Clocks A description of circadian rhythms in plants by de Mairan, Linnaeus, and Darwin
"Darkness Unveils Vital Metabolic Fuel Switch Between Sugar and Fat " (html) (in English). The
University of Texas Health Science Center (2006-01-18). Retrieved on 2007-10-17.
Circadian clocks go in vitro: purely post-translational oscillators in cyanobacteria
Journal of Biological Rhythms
Genetic switch for circadian rhythms discovered , University of California, Irvine, December 12,
2007.
CLOCK-mediated acetylation of BMAL1 controls circadian function , Nature (journal), 450, 1086-
1090 (13 December 2007).

Categories: Sleep | Circadian rhythms

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