Electromagnetic Fields and Circadian Rhythmicity (Circadian Factors in Human Health and Performance)
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At the molecular level, circadian modulation of memory could occur at multiple levels through regulation of sensory gating, neurotransmission, kinase signaling cascades, macromolecular synthesis, or epigenetic modifications. Circadian gating of sensory processes, as observed for olfaction in insects Krishnan et al. Downstream, molecular processes can serve as a bridge between the circadian clock and memory either through circadian regulation of basal expression and activity levels, rhythms in cellular compartmentalization, or rhythms in training-inducible activity or expression Table 1.
Neurotransmitters with circadian rhythms in basal expression and activity in brain regions associated with memory formation include acetylcholine, nitric oxide, serotonin, and GABA. Neurotransmitters can also function in conveyance of temporal information in memory context as suggested for acetylcholine in time-stamping Hut and Van der Zee and GABA synthesis in fear memory Bergado-Acosta et al.
An example of circadian regulation of training-induced neurotransmitter release comes from Aplysia , in which sensitization training-induced serotonin release is circadianly regulated with higher levels of serotonin observed in the hemolymph following training during the subjective day Lyons et al. Circadian regulation of inhibitory pathways may also modulate memory processes. In hamsters, circadian rhythmicity is necessary for declarative learning and memory as increased SCN GABAergic inhibition to the hippocampus in arrhythmic animals impairs memory Ruby et al.
Examples of molecular interactions between circadian systems and learning and memory. The phylogenetically conserved mechanisms of memory formation include intracellular second messenger and kinase signaling pathways.
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In the hippocampus, circadian cycling of phospho-ERK with peaks during the subjective day promotes memory formation and persistence of long-term memory Eckel-Mahan et al. Although a basal circadian rhythm in phospho-MAPK is not observed in Aplysia ganglia, sensitization training during the subjective day induces significantly greater MAPK activity compared with training during the night Lyons et al. Diurnal rhythms in cAMP levels, presumably due to adenylyl cyclase activity, occur in the hippocampus with peak levels during the day Eckel-Mahan et al.
Conversely, circadian regulation of memory may also occur through protein phosphatase activity. In Aplysia, inhibition of protein phosphatases rescues the induction of intermediate-term memory during the late subjective day and night Michel et al. Thus, the circadian regulation of intracellular signaling cascades through kinase and phosphatase regulation provides broad mechanisms through which the circadian clock can impact multiple types of learning and memory. The induction of gene expression, a necessary occurrence for LTM, provides multiple junctures through which the circadian clock can regulate memory formation.
More recently, hippocampal circadian rhythms in CREB activation have been shown to be dependent upon the Period1 gene Rawashdeh et al. In Drosophila mushroom bodies, a region necessary for many forms of memory, dCREB2 exhibits circadian oscillations in gene expression, nuclear protein levels and activity, likely mediating rhythms in LTM formation Tanenhaus et al.
Recent studies have highlighted the role of epigenetic modifications affecting gene transcription in the circadian regulation of memory formation. Acetylation of histone protein, particularly H3, enhances the induction and formation of long-term memory Levenson et al.
The circadian clock regulates H3 acetylation at lysine 14 in phase with trimethylation of H3 at lysine 4 providing a mechanism for circadian regulation of learning-induced gene transcription Rawashdeh et al. Thus, circadian regulation of learning-induced transcription necessary for LTM through epigenetic mechanisms or transcription factor activation provides a key mechanism through which the circadian clock targets memory formation.
The identification of modulated steps and targets of the circadian clock potentially may serve as the foundation for future therapies to improve memory and performance. Circadian modulation of human performance results in high economic costs to businesses attributable to industrial and occupational accidents, decreased worker productivity, and increased health costs for employees Rajaratnam et al.
Moreover, technological advances have resulted in increased human exposure to artificial light at night, particularly blue wavelengths that most affect the circadian clock, compounding the problems of circadian disorders in modern society for review, see Bonmati-Carrion et al. Consequently, identifying methods to maintain circadian rhythms or minimize circadian misalignment are necessary both at home and the workplace.
Recommendations by the American Academy of Sleep Medicine to ameliorate the problems of shift work are designed to minimize individual circadian misalignment and to increase alertness and performance Morgenthaler et al.
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In brief review, these strategies include: 1 the use of bright light therapy to enhance circadian re-entrainment Burgess et al. Readily passing through the blood brain barrier, melatonin increases daytime sleep duration with little effect on sleep latency for night-shift workers Sharkey et al. During the day, the circadian system maintains arousal in the face of increased time awake with decreased arousal at night due, in part, to rising melatonin levels which target MT 1 and MT 2 receptors in the SCN to dampen SCN driven arousal signals Dodson and Zee ; Carocci et al.
Melatonin is used as a treatment of circadian perturbations including jet lag, shift work, or delayed sleep disorder Pandi-Perumal et al. In forced desynchrony studies, repeated low doses of caffeine counteracted homeostatic sleep pressures, enhanced alertness, and ameliorated reductions in cognitive performance, particularly at the circadian troughs of performance Walsh et al. Modafinil and its R -enantiomer armodafinil improve attention and alertness, cognitive performance, long-term memory, and reduce sleepiness associated with night-shift work Walsh et al.
In addition to the above strategies designed to enable permanent shift workers to maintain regular rhythms minimizing circadian misalignment and enhancing performance, exercise during a night shift also may facilitate phase shifting and maintenance of circadian rhythms Eastman et al. For further review of strategies for managing shift work, see Boivin and James , Morgenthaler et al. For individuals engaged in rotating shift work given the length of time necessary for complete resynchronization of central and peripheral oscillators, reentrainment to the work cycle may not be optimal.
For these individuals, managing the direction of the rotation may reduce some negative health effects associated with shift work or even maintaining entrainment to a daytime light dark cycle through minimizing the effects of workplace lighting at night on melatonin secretion and hormonal rhythms. Recent research has suggested that the wearing of glasses with optical filters to block wavelengths shorter than or nm may attenuate circadian misalignment, improve mood disorders, enhance cognitive performance during shift work, and sleep upon cessation of shift work Rahman et al.
New applications and software also are becoming available to provide optical filters for personal electronic devices and home computers to minimize circadian perturbations from electronic devices during evening use.
Circadian Rhythm and Sleep Disruption: Causes, Metabolic Consequences, and Countermeasures
The realities of technological advancements in a modern, global society, and the risks to individual and public health due to circadian modulation of cognitive function or circadian desynchronization, emphasize the critical need for continued research to understand the molecular mechanisms underlying circadian and memory interactions, developing new therapeutic treatments and identifying practical measures to minimize individual circadian stresses. We thank Dr. Gregg Roman for helpful discussions and Miguel de la Flor for the brain illustration in Figure 1.
Research support for L. Synchrony and desynchrony in circadian clocks: impacts on learning and memory Harini C. Krishnan and Lisa C. Previous Section Next Section. Figure 1. View this table: In this window In a new window. Table 1. Previous Section.
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