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Circadian and Ultradian Clocks/Rhythmsq EW Lamont, Carleton University, Ottawa, ON, Canada S Amir, Concordia University, Montreal, QC, Canada

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Circadian and Ultradian Clocks/Rhythmsq EW Lamont, Carleton University, Ottawa, ON, Canada S Amir, Concordia University, Montreal, QC, Canada Circadian and Ultradian Clocks/Rhythmsq EW Lamont, Carleton University, Ottawa, ON, Canada S Amir, Concordia University, Montreal, QC, Canada Ó 2017 Elsevier Inc. All rights reserved. Introduction 1 Properties of Biological Clocks 2 Ultradian Rhythms 2 Circadian Rhythms 3 Molecular Mechanisms of Circadian Rhythms 3 Relationship of Circadian Rhythms to Physical and Mental Health 4 Relationship Between Ultradian and Circadian Clocks 4 Conclusion 5 References 5 Glossary Circadian biological rhythms of approximately 24 h in NREM non-rapid eye movement sleep, made up of several duration. sleep stages that are defined based on the frequency of Clock something that is used to measure time. brain electrical activity. Chronotherapeutics treatment protocol for cancer Oscillator a series of components with a combined action whereby the time of day that chemotherapy is that varies between extreme values over a precise period of administered is chosen in order to be maximally effective time. in treating the tumor, but have minimal effects on the Period the duration of a single oscillation measured from patient. peak to peak or from one reference point to the next ie, Entrainment the process by which an environmental from midnight to midnight on a 24-h clock. stimulus regulates the period and phase of an Phase the momentary state of an oscillation. oscillator. REM rapid eye movement sleep. Infradian rhythms with a period of days, weeks, or years. Ultradian rhythms with a period of less than 1 day. Introduction The term biological rhythm can be used to describe any molecular, physiological, or behavioral event or process that is recurring, and is a defining characteristic of all living organisms (Lloyd and Murray, 2005). Rhythmicity can be observed over the entire range of time scales from fractions of a second to years, but can be broadly grouped into three categories: ultradian, circadian, and infra- dian. Ultradian rhythms, such as cell division or the human rapid eye movement (REM)-non-REM (NREM) sleep cycle, can have a period, of seconds, minutes, or hours and occur multiple times per day. Circadian rhythms, including photosynthesis, the rest– activity cycle, and the plasma concentration of numerous circulating hormones, repeat approximately every 24-h. Infradian rhythms have periods of longer than 24 h, ranging from days to weeks, months, or even years. Examples include the human menstrual cycle, hibernation, and the 17-year emergence cycle of certain North American cicadas, respectively (see Fig. 1). The coordination of parallel and serial processes necessitates rhythmicity at all levels of biological organization. Ultradian rhythms likely arose in order to separate in time those mutually incompatible chemical reactions that could not be physically sepa- rated within the space of a single cell. In more complex systems, ultradian rhythmicity provides a metabolically inexpensive mech- anism for efficient signal transmission. For example, it may be energetically expensive or even toxic for an organism to have constantly high levels of a hormone in circulation, when the same signal can be transmitted by pulsatile hormone release (Lahav, 2004). Temporal separation of incompatible chemical reactions is also observed at a circadian time scale, such as the daily rhythm of photosynthesis. Circadian rhythmicity is also critical for the synchronization of multiple biological processes with the external environment and allows the organism to anticipate the environmental demands of the solar cycle. q Change History: January 2016. Elaine Waddington Lamont and Shimon Amir updated their affiliations, revised abstract, made some changes to the text, updated figures, replaced further reading with reference list and in-text citations. Reference Module in Neuroscience and Biobehavioral Psychology http://dx.doi.org/10.1016/B978-0-12-809324-5.00283-2 1 2 Circadian and Ultradian Clocks/Rhythms Figure 1 Time scale of biological rhythmicity, with examples. REM, Rapid eye movement; NREM, non rapid eye movement; GH, growth hormone; GnRH, gonadotropin-releasing hormone; LH, luteinizing hormone; FSH, follicle-stimulating hormone; TH, thyroid hormone. Properties of Biological Clocks In their most basic form, biological rhythms consist of a negative feedback loop with a time delay (Tiana et al., 2007). For example, there are numerous proteins that repress transcription of their own genes, such as the PERIOD (PER) and CRYPTOCHROME (CRY) proteins, essential components of the mammalian circadian clock (described in more detail below, and see Fig. 2). At least one negative feedback loop is necessary for an oscillation. Most biological clocks are made up of a series of weakly coupled oscillators, which make them more robust with respect to perturbations by outside stimuli. Greater complexity also tends to allow for timing of longer intervals. The most important characteristic, and hallmark of a true biological clock, is temperature compensation: the period of the oscil- lation must be very similar over a wide range of biological temperatures. This has been studied extensively in circadian rhythms, but is also a characteristic of ultradian clocks. Intrinsic rhythmicity and a free running period under constant conditions are also neces- sary properties for biological clocks. Ultradian Rhythms Because the time scale of ultradian rhythms is so broad, from fractions of a second to hours, a large number of biological clocks fall into this category and a comprehensive discussion is not possible in this entry. However, an example a of a well-studied ultradian rhythm is the approximately 40-min cycle of cellular respiration observed under continuous aerobic culture conditions in the yeast Figure 2 Molecular mechanisms of the mammalian circadian clock. Clock gene mRNAs are indicated by italics, proteins are in bold. Fine arrows indicate transcription, block arrows indicate binding or interaction, the gray arrow indicates translocation, and the open arrow indicates post- transcriptional modification. Excitation and inhibition are indicated by plus (þ) and minus (À) signs respectively. C, CLOCK protein; N, NPAS2 protein; B, BMAL1 protein. Circadian and Ultradian Clocks/Rhythms 3 Saccharomyces cerevisiae (Causton et al., 2015). Interestingly, the majority of the individual organisms within a culture will become synchronized with one another, such that the O2 concentration and intracellular pH of billions of cells become perfectly coordi- nated. This ultradian oscillator meets all of the requirements of a true clock. It is intrinsically rhythmic, temperature compensated, and maintains a 40-min oscillation for months under constant culture conditions (Causton et al., 2015). The signaling mechanism responsible for the mass inter-cellular synchronization has not yet been established, but two diffusible chemicals, H2S and acetal- dehyde, are possible candidates, as they show rhythmic changes in concentration and can shift the phase of the ultradian metabolic clock (Lloyd and Murray, 2005). The molecular mechanisms of this clock are under investigation and a putative clock genes include GTS1 (Ouyang et al., 2011; Wang et al., 2001), while PER-ARNT-SIM (PAS) domain containing PSK2 acts a nutrient sensing protein kinase that modulates the yeast ultradian clock (Ouyang et al., 2011). One of the first ultradian rhythms to be studied in humans was the REM-NREM sleep cycle, which has a period of about 90-min, and occurs 3–5 times in the average sleep episode (Moszczynski and Murray, 2012). This rhythm is composed of the synchronous activity of a number of different processes including occulomotor activity, muscle tone, dominance of the autonomic nervous system, brain electrical activity and energy utilization. Like many other biological processes, sleep shows rhythmicity at multiple time scales. In addition to a 90-min ultradian rhythm, a circadian rhythm is also evident (Dijk and Czeisler, 1995). Circadian Rhythms With the exception of extreme environments like the deep ocean floor, all organisms are exposed to the daily solar cycle and have an internal clock that cycles about once a day (Dunlap et al., 2004). For example, the unicellular marine algae Gonyaulax polyedra, moves up to the higher levels of the ocean to absorb sunlight and CO2 for photosynthesis during the day, but descends to the nutrient rich lower layers during the night to absorb nitrogen and phosphorous. In multicellular organisms, the molecular machinery for circadian rhythms is present in nearly every cell. These clocks perform functions specific to each individual tissue and are synchronized with the external environment by a master circadian pacemaker (Rosenwasser and Turek, 2015). In mammals, this master circadian clock is located deep in the brain in the suprachiasmatic nucleus (SCN) of the hypothalamus. The SCN has all of the defining attributes of a biological oscillator: intrinsic rhythmicity, a free running period under constant conditions, and temperature compensation. Substantial evidence indicates that the SCN is the master circa- dian clock, as SCN lesions abolish circadian rhythmicity of locomotor activity, feeding, drinking, and hormone release. Further- more, surgical transplantation of SCN cells can restore circadian rhythms of locomotor activity to recipient animals with SCN ablation. Hamsters with the tau mutation have an endogenous circadian rhythm with a period shorter than 24 h. Animals homo- zygous or heterozygous for the mutation have periods of 20 and 22 h respectively, whereas wild-type animals have
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