Theta wave - Knowledge

451:—and therefore has the potential to generate strong EEG signals. Basic EEG theory says that when a layer of neurons generates an EEG signal, the signal always phase-reverses at some level. Thus, theta waves recorded from sites above and below a generating layer have opposite signs. There are other complications as well: the hippocampal layers are strongly curved, and theta-modulated inputs impinge on them from multiple pathways, with varying phase relationships. The outcome of all these factors is that the phase and amplitude of theta oscillations change in a very complex way as a function of position within the hippocampus. The largest theta waves, however, are generally recorded from the vicinity of the fissure that separates the CA1 molecular layer from the dentate gyrus molecular layer. In rats, these signals frequently exceed 1 230:). In these species, the hippocampal oscillations fell mostly into the 4–6 Hz frequency range, so they were referred to as "theta" oscillations. Later, hippocampal oscillations of the same type were observed in rats; however, the frequency of rat hippocampal EEG oscillations averaged about 8 Hz and rarely fell below 6 Hz. Thus the rat hippocampal EEG oscillation should not, strictly speaking, have been called a "theta rhythm". However the term "theta" had already become so strongly associated with hippocampal oscillations that it continued to be used even for rats. Over the years this association has come to be stronger than the original association with a specific frequency range, but the original meaning also persists. 158:). This theory states that theta waves may act as a switch between encoding and recall mechanisms, and experimental data on rodents and humans support this idea. Another study on humans has showed that theta oscillations determine memory function (encoding or recall) when interacting with high frequency gamma activity in the hippocampus. These findings support the idea that theta oscillations support memory formation and retrieval in interaction with other oscillatory rhythms. These different theories have since been combined, as it has been shown that the firing patterns can support both navigation and memory. 317:, has a frequency in the 4–7 Hz range, and is eliminated by administration of atropine. Many later investigations have supported the general concept that hippocampal theta can be divided into two types, although there has been dispute about the precise properties of each type. Type 2 theta is comparatively rare in unanesthetized rats: it may be seen briefly when an animal is preparing to make a movement but hasn't yet executed it, but has only been reported for extended periods in animals that are in a state of frozen immobility because of the nearby presence of a predator such as a cat or ferret ( 285:. The frequency of the theta waves increases as a function of running speed, starting at about 6.5 Hz on the low end, and increasing to about 9 Hz at the fastest running speeds, although higher frequencies are sometimes seen for brief high-velocity movements such as jumps across wide gaps. In larger species of animals, theta frequencies are generally lower. The behavioral dependency also seems to vary by species: in cats and rabbits, theta is often observed during states of motionless alertness. This has been reported for rats as well, but only when they are fearful ( 566:) found that oscillations in the 4–7 Hz frequency range could be recorded from both the hippocampus and neocortex. The hippocampal oscillations were associated with REM sleep and the transition from sleep to waking, and came in brief bursts, usually less than a second long. Cortical theta oscillations were observed during the transition from sleep and during quiet wakefulness; however, the authors were unable to find any correlation between hippocampal and cortical theta waves, and concluded that the two processes are probably controlled by independent mechanisms. 333:) made a strong argument that the presence of theta in the hippocampal EEG can be predicted on the basis of what an animal is doing, rather than why the animal is doing it. Active movements such as running, jumping, bar-pressing, or exploratory sniffing are reliably associated with theta; inactive states such as eating or grooming are associated with LIA. Later studies showed that theta frequently begins several hundred milliseconds before the onset of movement, and that it is associated with the intention to move rather than with feedback produced by movement ( 542:

number of EEG studies have been conducted using electrodes glued to the scalp. The signals picked up by scalp electrodes are comparatively small and diffuse and arise almost entirely from the cerebral cortex for the hippocampus is too small and too deeply buried to generate recognizable scalp EEG signals. Human EEG recordings show clear theta rhythmicity in some situations, but because of the technical difficulties, it has been difficult to tell whether these signals have any relationship with the hippocampal theta signals recorded from other species.

466:). The dentate gyrus also generates theta waves, which are difficult to separate from the CA1 waves because they are considerably smaller in amplitude, but there is some evidence that dentate gyrus theta is usually about 90 degrees out of phase from CA1 theta. Direct projections from the septal area to hippocampal interneurons also play a role in generating theta waves, but their influence is much smaller than that of the entorhinal inputs (which are, however, themselves controlled by the septum). 186:). Their findings provoked widespread interest, in part because they related hippocampal activity to arousal, which was at that time the hottest topic in neuroscience. Green and Arduini described an inverse relationship between hippocampal and cortical activity patterns, with hippocampal rhythmicity occurring alongside desynchronized activity in the cortex, whereas an irregular hippocampal activity pattern was correlated with the appearance of large slow waves in the cortical EEG. 525:(LTP) to reinforce the connections between neurons of the hippocampus representing subsequent elements of a memory sequence. Indeed, it has been suggested that stimulation at the theta frequency is optimal for the induction of hippocampal LTP. Based on evidence from electrophysiological studies showing that both synaptic plasticity and strength of inputs to hippocampal region CA1 vary systematically with ongoing theta oscillations ( 154:, and proposed that it is related to arousal. Vanderwolf and his colleagues, noting the strong relationship between theta and motor behavior, have argued that it is related to sensorimotor processing. Another school, led by John O'Keefe, have suggested that theta is part of the mechanism animals use to keep track of their location within the environment. Another theory links the theta rhythm to mechanisms of learning and memory ( 258: 41: 376:). Some of the projections from the medial septal area are cholinergic; the rest are GABAergic or glutamatergic. It is commonly argued that cholinergic receptors do not respond rapidly enough to be involved in generating theta waves, and therefore that GABAergic and/or glutamatergic signals (Ujfalussy and Kiss, 2006) must play the central role. 349:). Lesioning the medial septal area, or inactivating it with drugs, eliminates both type 1 and type 2 theta. Under certain conditions, theta-like oscillations can be induced in hippocampal or entorhinal cells in the absence of septal input, but this does not occur in intact, undrugged adult rats. The critical septal region includes the 221:

literature dating back to the 1920s, Greek letters such as alpha, beta, theta, and gamma were used to classify EEG waves falling into specific frequency ranges, with "theta" generally meaning a range of about 4–7 cycles per second (Hz). In the 1930s–1950s, a very strong rhythmic oscillation pattern

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In humans, hippocampal theta rhythm has been observed and linked to memory formation and navigation. As with rats, humans exhibit hippocampal theta wave activity during REM sleep. Humans also exhibit predominantly cortical theta wave activity during REM sleep. Increased sleepiness is associated with

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A major research problem has been to discover the "pacemaker" for the theta rhythm, that is, the mechanism that determines the oscillation frequency. The answer is not yet entirely clear, but there is some evidence that type 1 and type 2 theta depend on different pacemakers. For type 2 theta, the

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In non-human animals, EEG signals are usually recorded using electrodes implanted in the brain; the majority of theta studies have involved electrodes implanted in the hippocampus. In humans, because invasive studies are not ethically permissible except in some neurological patients, the largest

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Theta is not just confined to the hippocampus. In rats, it can be observed in many parts of the brain, including nearly all that interact strongly with the hippocampus. The generation of the rhythm is dependent on the medial septal area: this area projects to all of the regions that show theta

1536:"The remarkable thing, however, is that as the meditators signalled that they had entered into the state of mental silence, or “thoughtless awareness”, another form of brain wave activity emerged which involved “theta waves” focused specifically in the front and top of the brain in the midline." 1219:

Saint Amour di Chanaz L, Pérez-Bellido A, Wu X, Lozano-Soldevilla D, Pacheco-Estefan D, Lehongre K, Conde-Blanco E, Roldan P, Adam C, Lambrecq V, Frazzini V, Donaire A, Carreño M, Navarro V, Valero-Cabré A, Fuentemilla L (May 2023). "Gamma amplitude is coupled to opposed hippocampal theta-phase

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and several brainstem areas. Hippocampal theta rhythms in other species differ in some respects from those in rats. In cats and rabbits, the frequency range is lower (around 4–6 Hz), and theta is less strongly associated with movement than in rats. In bats, theta appears in short bursts

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along the theta wave in the hippocampus permits neural signals representing events that are only expected or those from the recent past to be placed next to the actually ongoing ones along a single theta cycle, and to be repeated over several theta cycles. This mechanism is supposed to allow

182:). They were not able to follow up on these initial observations, and it was not until 1954 that further information became available, in a very thorough study by John D. Green and Arnaldo Arduini that mapped out the basic properties of hippocampal oscillations in cats, rabbits, and monkeys ( 198:). A major contribution came from a group of investigators working in Vienna, including Stumpf and Wolfgang Petsche, who established the critical role of the medial septum in controlling hippocampal electrical activity, and worked out some of the pathways by which it exerts its influence. 337:). The faster an animal runs, the higher the theta frequency. In rats, the slowest movements give rise to frequencies around 6.5 Hz, the fastest to frequencies around 9 Hz, although faster oscillations can be observed briefly during very vigorous movements such as large jumps. 244:

The first meaning is usually intended in literature that deals with rats or mice, while the second meaning is usually intended in studies of human EEG recorded using electrodes glued to the scalp. In general, it is not safe to assume that observations of "theta" in the human EEG have any

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refers to frequency components in the 4–7 Hz range, regardless of their source. Cortical theta is observed frequently in young children. In older children and adults, it tends to appear during meditative, drowsy, hypnotic or sleeping states, but not during the deepest stages of

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Due to the density of its neural layers, the hippocampus generates some of the largest EEG signals of any brain structure. In some situations the EEG is dominated by regular waves at 4–10 Hz, often continuing for many seconds. This EEG pattern is known as the

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Over the following decade came an outpouring of experiments examining the pharmacology and physiology of theta. By 1965, Charles Stumpf was able to write a lengthy review of "Drug action on the electrical activity of the hippocampus" citing hundreds of publications

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In contrast to the situation in rats, where long periods of theta oscillations are easily observed using electrodes implanted at many sites, theta has been difficult to pin down in primates, even when intracortical electrodes have been available. Green and Arduini

533:), it has been suggested that the theta rhythm functions to separate periods of encoding of current sensory stimuli and retrieval of episodic memory cued by current stimuli so as to avoid interference that would occur if encoding and retrieval were simultaneous. 557:

Most of the available information on human hippocampal theta comes from a few small studies of epileptic patients with intracranially implanted electrodes used as part of a treatment plan. In the largest and most systematic of these studies, Cantero

114:. Theta waves with a lower frequency range, usually around 6–7 Hz, are sometimes observed when a rat is motionless but alert. When a rat is eating, grooming, or sleeping, the hippocampal EEG usually shows a non-rhythmic pattern known as 309:). Type 1 ("atropine resistant") theta, according to them, appears during locomotion and other types of "voluntary" behavior and during REM sleep, has a frequency usually around 8 Hz, and is unaffected by the anticholinergic drug 149:

The function of the hippocampal theta rhythm is not clearly understood. Green and Arduini, in the first major study of this phenomenon, noted that hippocampal theta usually occurs together with desynchronized EEG in the

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Aftanas, LI; Golocheikine, SA (September 2001). "Human anterior and frontal midline theta and lower alpha reflect emotionally positive state and internalized attention: high-resolution EEG investigation of meditation".

462:, via the direct EC→CA1 pathway. Another important driving force comes from the CA3→CA1 projection, which is out of phase with the entorhinal input, leading to a gradual phase shift as a function of depth within CA1 ( 2142:

Kramis R, Vanderwolf CH, Bland BH (1975). "Two types of hippocampal rhythmical slow activity in both the rabbit and the rat: relations to behavior and effects of atropine, diethyl ether, urethane, and pentobarbital".

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Several types of hippocampal and entorhinal neurons are capable of generating theta-frequency membrane potential oscillations when stimulated. Typically these are sodium-dependent voltage-sensitive oscillations in

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In rats, theta wave rhythmicity is easily observed in the hippocampus, but can also be detected in numerous other cortical and subcortical brain structures. Hippocampal theta waves, with a frequency range of

384:). For type 1 theta, the picture is still unclear, but the most widely accepted hypothesis proposes that the frequency is determined by a feedback loop involving the medial septal area and hippocampus ( 2447:

Whishaw IQ, Vanderwolf CH (1973). "Hippocampal EEG and behavior: changes in amplitude and frequency of RSA (theta rhythm) associated with spontaneous and learned movement patterns in rats and cats".

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As a rule, EEG signals are generated by synchronized synaptic input to the dendrites of neurons arranged in a layer. The hippocampus contains multiple layers of very densely packed neurons—the

554:) described a clear 7–9 Hz theta rhythm in the hippocampus of urethane-anesthetized macaques and squirrel monkeys, resembling the type 2 theta observed in urethane-anesthetized rats. 170:. Theta from the midfrontal cortex is specifically related to cognitive control and alterations in these theta signals are found in multiple psychiatric and neurodevelopmental disorders. 360:

The medial septal area projects to a large number of brain regions that show theta modulation, including all parts of the hippocampus as well as the entorhinal cortex, perirhinal cortex,

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In rats, hippocampal theta is seen mainly in two conditions: first, when an animal is running, walking, or in some other way actively interacting with its surroundings; second, during

550:), in their pioneering study of theta rhythms, reported only brief bursts of irregular theta in monkeys. Other investigators have reported similar results, although Stewart and Fox ( 178:

Although there were a few earlier hints, the first clear description of regular slow oscillations in the hippocampal EEG came from a paper written in German by Jung and Kornmüller (

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Kirk IJ (1998). "Frequency modulation of hippocampal theta by the supramammillary nucleus, and other hypothalamo-hippocampal interactions: mechanisms and functional implications".

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In 1975 Kramis, Bland, and Vanderwolf proposed that in rats there are two distinct types of hippocampal theta rhythm, with different behavioral and pharmacological properties (

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Bland, BH; Oddie SD (2001). "Theta band oscillation and synchrony in the hippocampal formation and associated structures: the case for its role in sensorimotor integration".

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Brankack, J; Stewart M; Fox SE (1993). "Current source density analysis of the hippocampal theta rhythm: Associated sustained potentials and candidate synaptic generators".

194:), and in 1964 John Green, who served as the leader of the field during this period, was able to write an extensive and detailed review of hippocampal electrophysiology ( 3214: 1919:

Hasselmo, ME (2005). "What is the Function of Hippocampal Theta Rhythm?- Linking Behavioral Data to Phasic Properties of Field Potential and Unit Recording Data".

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Larson, J.; Wong, D.; Lynch, G. (1986-03-19). "Patterned stimulation at the theta frequency is optimal for the induction of hippocampal long-term potentiation".

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that underlies various aspects of cognition and behavior, including learning, memory, and spatial navigation in many animals. It can be recorded using various

3237: 2825: 249:, and even if it falls into a certain frequency range, this cannot be taken to indicate that it has any functional dependence on the hippocampus. 967: 482:). Conversely, theta oscillations have been correlated to various voluntary behaviors (exploration, spatial navigation, etc.) and alert states ( 2349: 769: 2015:"Stimulation in hippocampal region CA1 in behaving rats yields LTP when delivered to the peak of theta and LTD when delivered to the trough" 3468: 3207: 99: 2787: 240:

EEG oscillations in the 4–7 Hz frequency range, regardless of where in the brain they occur or what their functional significance is.

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in amplitude. Theta waves recorded from above the hippocampus are smaller, and polarity-reversed with respect to the fissure signals.

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Buzsáki, G (2005). "Theta rhythm of navigation: link between path integration and landmark navigation, episodic and semantic memory".

357:. The lateral septal nucleus, a major recipient of hippocampal output, probably does not play an essential role in generating theta. 1439: 478:). Studies suggest that these rhythms reflect the "on-line" state of the hippocampus; one of readiness to process incoming signals ( 2665: 2660: 2632: 2183:

Jung, R; Kornmüller AE (1938). "Eine Methodik der ableitung lokalisierter Potentialschwankungen aus subcorticalen Hirngebieten".

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Alonso, A; Llinás R (1989). "Subthreshold Na-dependent theta-like rhythmicity in entorhinal cortex layer II stellate cells".

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Wang XJ (2002). "Pacemaker neurons for the theta rhythm and their synchronization in the septohippocampal reciprocal loop".

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The strongest theta waves are generated by the CA1 layer, and the most significant input driving them comes from the

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of the hypothalamus, anterior nuclei of the thalamus, amygdala, inferior colliculus, and several brainstem nuclei (

2715: 2983: 2792: 2740: 1072: 110:, appear when a rat is engaged in active motor behavior such as walking or exploratory sniffing, and also during 3494: 3453: 3338: 2540: 1537: 494:). A large body of evidence indicates that theta rhythm is likely involved in spatial learning and navigation ( 282: 275: 267: 237:

A specific type of regular oscillation seen in the hippocampus and several other brain regions connected to it.

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are low-frequency components of scalp EEG, usually recorded from humans. Theta rhythms can be quantified using

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Numerous studies have shown that the medial septal area plays a central role in generating hippocampal theta (

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Sainsbury, RS; Heynen A; Montoya CP (1987). "Behavioral correlates of hippocampal type 2 theta in the rat".

1019: 490:), suggesting that it may reflect the integration of sensory information with motor output (for review, see 369: 656:

Seager, Matthew A.; Johnson, Lynn D.; Chabot, Elizabeth S.; Asaka, Yukiko; Berry, Stephen D. (2002-02-05).

214:. Both of these are oscillatory EEG patterns, but they may have little in common beyond the name "theta". 3283: 3252: 3223: 3176: 2725: 2560: 1928: 1766: 1355:"Midfrontal Theta Activity in Psychiatric Illness: An Index of Cognitive Vulnerabilities Across Disorders" 914: 715:

Winson, J. (1978-07-14). "Loss of hippocampal theta rhythm results in spatial memory deficit in the rat".

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Theta-frequency activity arising from the hippocampus is manifested during some short-term memory tasks (

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Because of a historical accident, the term "theta rhythm" is used to refer to two different phenomena,

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Vanderwolf, C. H (1 April 1969). "Hippocampal electrical activity and voluntary movement in the rat".

3247: 3152: 1552: 857: 724: 669: 361: 1933: 1771: 919: 3427: 3368: 3353: 3328: 3318: 3308: 3147: 3137: 2603: 2503: 1261: 1015:"Decreased electrophysiological activity represents the conscious state of emptiness in meditation" 623: 313:. Type 2 ("atropine sensitive") theta appears during immobility and during anesthesia induced by 3448: 3384: 3343: 3157: 2910: 2810: 2675: 2433: 2394: 2318: 2279: 2240: 2200: 2168: 2128: 1954: 1907: 1792: 1743: 1698: 1659: 1620: 1577: 1519: 1476: 1335: 992: 940: 826: 444: 409: 393: 56: 573:

with stronger theta-frequency activity as well as with changes to the gamma-frequency activity (

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relationship to the "hippocampal theta rhythm". Scalp EEG is generated almost entirely by the

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cortex during learning and memory retrieval, and are believed to be vital to the induction of

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The sample of mouse EEG. Theta rhythm is prominent during part of awaking and REM sleep.

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Ekstrom, Arne D. (2005). "Human hippocampal theta activity during virtual navigation".

71:(EEG), recorded either from inside the brain or from electrodes attached to the scalp. 2460: 2341: 2116: 1730: 1713: 1647: 1565: 692: 657: 3483: 3422: 3085: 2933: 2890: 2840: 2835: 2608: 2306: 2267: 2228: 2156: 1686: 1507: 1480: 1411: 880: 845: 440: 413: 123: 2398: 2322: 2283: 2204: 2132: 1911: 1796: 1747: 1702: 1663: 1581: 1523: 1339: 980: 846:"Theta oscillations index human hippocampal activation during a working memory task" 830: 3412: 3053: 2830: 2815: 2437: 2172: 1958: 1624: 1373: 1354: 996: 944: 257: 127: 2254:

Stewart M, Fox SE (1990). "Do septal neurons pace the hippocampal theta rhythm?".

2244: 1895: 1307: 2075: 1985: 1323: 1170:"Phase separation of competing memories along the human hippocampal theta rhythm" 1136: 1121:"Gradual changes in hippocampal activity support remembering the order of events" 3112: 2988: 2928: 2880: 2875: 2866: 2705: 2655: 2650: 2532: 505: 483: 223: 79: 75: 3192: 2332:

Stumpf, C (1965). "Drug action on the electrical activity of the hippocampus".

960:"A systematic review of the neurophysiology of mindfulness on EEG oscillations" 662:

Proceedings of the National Academy of Sciences of the United States of America

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Vertes, RP (2005). "Hippocampal theta rhythm: a tag for short-term memory".

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http://www.researchingmeditation.org/meditation-research-summary/brain-waves

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McLoughlin, Gráinne; Gyurkovics, Máté; Palmer, Jason; Makeig, Scott (2021).

629: 448: 278:(LIA) that usually dominates the hippocampal EEG when theta is not present. 151: 2429: 2390: 2093: 2048: 2003: 1950: 1903: 1872: 1841: 1808:"Sleep-dependent theta oscillations in the human hippocampus and neocortex" 1806:

Cantero JL, Atienza M, Stickgold R, Kahana MJ, Madsen JR, Kocsis B (2003).

1788: 1739: 1655: 1573: 1472: 1382: 1331: 1292: 1241: 1205: 1154: 1105: 1052: 988: 936: 889: 822: 736: 701: 682: 122:. The hippocampal theta rhythm depends critically on projections from the 17: 2468: 2421: 2359: 2314: 2275: 2236: 2164: 2124: 1694: 1616: 1515: 1419: 1257:"Memory, navigation and theta rhythm in the hippocampal-entorhinal system" 1220:

states during the encoding and retrieval of episodic memories in humans".

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rhythmicity, and destruction of it eliminates theta throughout the brain (

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Hyman, JM; Wyble BP; Goyal V; Rossi CA; Hasselmo ME (December 17, 2003).

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supramammillary nucleus of the hypothalamus appears to exert control (

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Stewart M, Fox SE (1991). "Hippocampal theta activity in monkeys".

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Green, JD; Arduini A (1954). "Hippocampal activity in arousal".

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Kerrén C, van Bree S, Griffiths BJ, Wimber M (November 2022).

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OREKHOVA, E; STROGANOVA, T; POSIKERA, I; ELAM, M (May 2006).

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At least two types of theta rhythm have been described. The

577:). Also, increased theta waves have been seen in humans in ' 107: 1066:

Lee DJ, Kulubya E, Goldin P, Goodarzi A, Girgis F (2018).

1068:"Review of the Neural Oscillations Underlying Meditation" 516:, a potential cellular mechanism of learning and memory. 86:

including rodents, rabbits, dogs, cats, and marsupials.

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Hinterberger T, Schmidt S, Kamei T, Walach H (2014).

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is a strong oscillation that can be observed in the

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Electroencephalography and Clinical Neurophysiology

1119:Manns JR, Howard MW, Eichenbaum H (November 2007). 3238:Amplitude integrated electroencephalography (aEEG) 82:and other brain structures in numerous species of 416:, these oscillations result from an interplay of 850:Proceedings of the National Academy of Sciences 1008: 1006: 3208: 2504: 233:Thus, "theta" can mean either of two things: 8: 420:excitation via a persistent sodium current ( 94:using freely available toolboxes, such as, 3469:Neurophysiological Biomarker Toolbox (NBT) 3215: 3201: 3193: 2826:Rapid eye movement sleep behavior disorder 2628: 2511: 2497: 2489: 1255:Buzsáki, György; Moser, Edvard I. (2013). 92:quantitative electroencephalography (qEEG) 2083: 2064:The American Journal of Clinical Hypnosis 2038: 1993: 1932: 1831: 1770: 1729: 1462: 1452: 1372: 1282: 1195: 1185: 1144: 1095: 1085: 1042: 1032: 918: 879: 869: 812: 691: 681: 2484:Brain slice models of theta EEG activity 2058:Jensen MP, Adachi T, Hakimian S (2015). 968:Neuroscience & Biobehavioral Reviews 146:has been shown to increase theta power. 126:, which in turn receives input from the 1714:"Theta oscillations in the hippocampus" 844:Tesche, C. D.; Karhu, J. (2000-01-18). 648: 487: 330: 1882:Green, JD (1964). "The hippocampus". 758:Squire, Larry R. (17 December 2012). 569:Studies have shown an association of 318: 286: 155: 30:"Thetawaves" redirects here. For the 7: 2334:International Review of Neurobiology 530: 463: 404:). Specifically, it appears that in 100:Neurophysiological Biomarker Toolbox 3324:Contingent negative variation (CNV) 3263:Brainstem auditory evoked potential 1968:Hasselmo, ME; Eichenbaum H (2005). 563: 551: 495: 479: 428: 373: 346: 334: 294: 2031:10.1523/JNEUROSCI.23-37-11725.2003 1824:10.1523/JNEUROSCI.23-34-10897.2003 574: 401: 306: 191: 25: 1440:Frontiers in Systems Neuroscience 958:Lomas T, Ivtzan I, Fu CH (2015). 547: 526: 501:Theta rhythms are very strong in 491: 227: 195: 183: 2666:Obesity hypoventilation syndrome 2661:Central hypoventilation syndrome 2336:. Vol. 8. pp. 77–138. 385: 381: 179: 142:and increased theta wave power. 2821:Periodic limb movement disorder 2788:Non-24-hour sleep–wake disorder 981:10.1016/j.neubiorev.2015.09.018 427:) with perisomatic inhibition ( 274:(RSA), to contrast it with the 161:In human EEG studies, the term 3258:Somatosensory evoked potential 1374:10.1016/j.biopsych.2021.08.020 475: 335:Whishaw & Vanderwolf, 1973 131:associated with echolocation. 1: 3454:Difference due to memory (Dm) 3093:Biphasic and polyphasic sleep 2901:Nocturnal clitoral tumescence 2763:Advanced sleep phase disorder 2461:10.1016/S0091-6773(73)80041-0 2342:10.1016/S0074-7742(08)60756-4 2117:10.1016/S0149-7634(97)00015-8 1896:10.1152/physrev.1964.44.4.561 1731:10.1016/S0896-6273(02)00586-X 1648:10.1016/S0166-4328(01)00358-8 1566:10.1016/S0304-3940(01)02094-8 353:and the vertical limb of the 3253:Magnetoencephalography (MEG) 3224:Electroencephalography (EEG) 2773:Delayed sleep phase disorder 2701:Excessive daytime sleepiness 2307:10.1016/0006-8993(91)90376-7 2268:10.1016/0166-2236(90)90040-H 2229:10.1016/0031-9384(87)90382-9 2157:10.1016/0014-4886(75)90195-8 2076:10.1080/00029157.2014.976786 1986:10.1016/j.neunet.2005.08.007 1687:10.1016/0006-8993(93)90043-M 1508:10.1016/0006-8993(86)90579-2 1433:Kovács KA (September 2020). 1412:10.1016/0013-4694(69)90092-3 1324:10.1016/j.clinph.2005.12.027 1137:10.1016/j.neuron.2007.08.017 3248:Electrocorticography (ECoG) 2906:Nocturnal penile tumescence 2778:Irregular sleep–wake rhythm 270:. It has also been called 3521: 2768:Cyclic alternating pattern 608:Theta wave — (4–8 Hz) 325:Relationship with behavior 29: 27:Neural oscillatory pattern 2984:Behavioral sleep medicine 2793:Shift work sleep disorder 2741:Sleep state misperception 1454:10.3389/fnsys.2020.559186 1234:10.1016/j.cub.2023.03.073 1073:Frontiers in Neuroscience 793:Lega, Bradley C. (2011). 537:Humans and other primates 402:Alonso & Llinás, 1989 228:Green & Arduini, 1954 2541:Rapid eye movement (REM) 1865:10.1152/jn.1954.17.6.533 1312:Clinical Neurophysiology 1087:10.3389/fnins.2018.00178 1034:10.3389/fpsyg.2014.00099 761:Fundamental neuroscience 276:large irregular activity 268:hippocampal theta rhythm 116:large irregular activity 88:"Cortical theta rhythms" 76:hippocampal theta rhythm 3375:Late positive component 3243:Event-related potential 2862:Exploding head syndrome 2671:Obstructive sleep apnea 1020:Frontiers in Psychology 492:Bland & Oddie, 2001 347:Stewart & Fox, 1990 295:Stewart & Fox, 1990 184:Green and Arduini, 1954 3490:Electroencephalography 3284:Bereitschaftspotential 3177:Sleeping while on duty 2726:Idiopathic hypersomnia 2185:Arch Psychiat Nervenkr 737:10.1126/science.663646 683:10.1073/pnas.032662099 579:no thought' meditation 523:long term potentiation 514:long-term potentiation 370:supramammillary nuclei 355:diagonal band of Broca 319:Sainsbury et al., 1987 287:Sainsbury et al., 1987 272:Rhythmic Slow Activity 262: 222:was discovered in the 212:"human cortical theta" 45: 2999:Neuroscience of sleep 2731:Night eating syndrome 2716:Kleine–Levin syndrome 2422:10.1152/jn.00135.2001 2105:Neurosci Biobehav Rev 1360:Biological Psychiatry 871:10.1073/pnas.97.2.919 626:— (12.5–15.5 Hz) 531:Brankack et al., 1993 464:Brankack, et al. 1993 351:medial septal nucleus 260: 226:of cats and rabbits ( 43: 3153:Sleep and creativity 1553:Neuroscience Letters 620:— (7.5–12.5 Hz) 362:retrosplenial cortex 69:electroencephalogram 65:electrophysiological 3428:Sensorimotor rhythm 3385:Neural oscillations 3329:Mismatch negativity 3148:Sleep and breathing 2604:Sensorimotor rhythm 1712:Buzsáki, G (2002). 1262:Nature Neuroscience 1187:10.7554/eLife.80633 862:2000PNAS...97..919T 729:1978Sci...201..160W 674:2002PNAS...99.1616S 575:Jensen et al., 2015 307:Kramis et al., 1975 208:"hippocampal theta" 3158:Sleep and learning 2911:Nocturnal emission 2811:Nightmare disorder 2676:Periodic breathing 2383:10.1002/hipo.20118 2197:10.1007/BF02157817 1943:10.1002/hipo.20116 1781:10.1002/hipo.20113 929:10.1002/hipo.20109 814:10.1002/hipo.20937 638:— (32–100 Hz) 527:Hyman et al., 2003 394:membrane potential 263: 124:medial septal area 57:neural oscillation 46: 3500:Electrophysiology 3477: 3476: 3371:(late positivity) 3273:Evoked potentials 3190: 3189: 3168:Sleep deprivation 3007: 3006: 2351:978-0-12-366808-0 2151:(1 Pt 1): 58–85. 1818:(34): 10897–903. 1228:(9): 1836–43.e6. 771:978-0-12-385871-9 632:— (16–31 Hz) 605:— (0.1–3 Hz) 486:, etc.) in rats ( 470:Research findings 460:entorhinal cortex 301:Type 1 and type 2 67:methods, such as 44:An EEG theta wave 36:Steal This Album! 16:(Redirected from 3512: 3459:Oddball paradigm 3217: 3210: 3203: 3194: 3163:Sleep and memory 3103:Circadian rhythm 2850:Benign phenomena 2752:Circadian rhythm 2629: 2513: 2506: 2499: 2490: 2472: 2441: 2402: 2363: 2326: 2287: 2248: 2209: 2208: 2176: 2136: 2097: 2087: 2052: 2042: 2025:(37): 11725–31. 2007: 1997: 1962: 1936: 1915: 1876: 1845: 1835: 1800: 1774: 1751: 1733: 1706: 1667: 1628: 1609:10.1038/342175a0 1586: 1585: 1546: 1540: 1534: 1528: 1527: 1491: 1485: 1484: 1466: 1456: 1430: 1424: 1423: 1393: 1387: 1386: 1376: 1350: 1344: 1343: 1303: 1297: 1296: 1286: 1252: 1246: 1245: 1216: 1210: 1209: 1199: 1189: 1165: 1159: 1158: 1148: 1116: 1110: 1109: 1099: 1089: 1063: 1057: 1056: 1046: 1036: 1010: 1001: 1000: 964: 955: 949: 948: 922: 900: 894: 893: 883: 873: 841: 835: 834: 816: 790: 784: 783: 755: 749: 748: 712: 706: 705: 695: 685: 653: 614:— (8–15 Hz) 518:Phase precession 488:Vanderwolf, 1969 398:action potential 32:System of a Down 21: 3520: 3519: 3515: 3514: 3513: 3511: 3510: 3509: 3495:Neurophysiology 3480: 3479: 3478: 3473: 3437: 3379: 3267: 3226: 3221: 3191: 3186: 3081:Procrastination 3034:Four-poster bed 3003: 2967: 2961:Polysomnography 2939:Sleep induction 2915: 2886:Sleep paralysis 2845: 2797: 2756: 2753: 2745: 2687: 2646:Mouth breathing 2624:Sleep disorders 2618: 2555: 2546:Quiescent sleep 2526: 2524:sleep disorders 2517: 2480: 2475: 2446: 2407: 2368: 2352: 2331: 2292: 2256:Trends Neurosci 2253: 2214: 2182: 2181: 2141: 2102: 2057: 2012: 1974:Neural Networks 1967: 1934:10.1.1.483.5818 1918: 1881: 1850: 1805: 1772:10.1.1.476.6199 1756: 1711: 1672: 1642:(1–2): 119–36. 1636:Behav Brain Res 1633: 1603:(6246): 175–7. 1594: 1590: 1589: 1548: 1547: 1543: 1535: 1531: 1493: 1492: 1488: 1432: 1431: 1427: 1395: 1394: 1390: 1352: 1351: 1347: 1305: 1304: 1300: 1275:10.1038/nn.3304 1254: 1253: 1249: 1218: 1217: 1213: 1167: 1166: 1162: 1118: 1117: 1113: 1065: 1064: 1060: 1012: 1011: 1004: 962: 957: 956: 952: 920:10.1.1.535.1693 902: 901: 897: 843: 842: 838: 792: 791: 787: 772: 757: 756: 752: 723:(4351): 160–3. 714: 713: 709: 655: 654: 650: 645: 599: 587: 539: 472: 449:subicular layer 437: 426: 343: 327: 303: 255: 247:cerebral cortex 204: 176: 38: 28: 23: 22: 15: 12: 11: 5: 3518: 3516: 3508: 3507: 3502: 3497: 3492: 3482: 3481: 3475: 3474: 3472: 3471: 3466: 3461: 3456: 3451: 3445: 3443: 3439: 3438: 3436: 3435: 3430: 3425: 3420: 3415: 3410: 3405: 3400: 3395: 3389: 3387: 3381: 3380: 3378: 3377: 3372: 3366: 3361: 3356: 3351: 3346: 3341: 3336: 3332: 3331: 3326: 3321: 3316: 3311: 3306: 3301: 3296: 3291: 3286: 3281: 3277: 3275: 3269: 3268: 3266: 3265: 3260: 3255: 3250: 3245: 3240: 3234: 3232: 3228: 3227: 3222: 3220: 3219: 3212: 3205: 3197: 3188: 3187: 3185: 3184: 3179: 3174: 3165: 3160: 3155: 3150: 3145: 3140: 3135: 3130: 3125: 3120: 3115: 3110: 3108:Comfort object 3105: 3100: 3095: 3090: 3089: 3088: 3083: 3073: 3068: 3063: 3058: 3057: 3056: 3051: 3046: 3041: 3036: 3031: 3026: 3015: 3013: 3009: 3008: 3005: 3004: 3002: 3001: 2996: 2991: 2986: 2981: 2979:Sleep medicine 2975: 2973: 2969: 2968: 2966: 2965: 2964: 2963: 2953: 2952: 2951: 2946: 2936: 2931: 2925: 2923: 2917: 2916: 2914: 2913: 2908: 2903: 2898: 2893: 2888: 2883: 2878: 2869: 2864: 2859: 2853: 2851: 2847: 2846: 2844: 2843: 2838: 2833: 2828: 2823: 2818: 2813: 2807: 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1367:(2): 173–182. 1345: 1318:(5): 1047–62. 1298: 1247: 1211: 1160: 1111: 1058: 1002: 950: 913:(7): 881–889. 895: 856:(2): 919–924. 836: 807:(4): 748–761. 785: 770: 750: 707: 668:(3): 1616–20. 647: 646: 644: 641: 640: 639: 633: 627: 621: 615: 609: 606: 598: 595: 594: 593: 586: 583: 538: 535: 471: 468: 436: 433: 424: 342: 339: 326: 323: 302: 299: 254: 251: 242: 241: 238: 217:In the oldest 203: 200: 175: 172: 156:Hasselmo, 2005 26: 24: 14: 13: 10: 9: 6: 4: 3: 2: 3517: 3506: 3503: 3501: 3498: 3496: 3493: 3491: 3488: 3487: 3485: 3470: 3467: 3465: 3462: 3460: 3457: 3455: 3452: 3450: 3447: 3446: 3444: 3440: 3434: 3431: 3429: 3426: 3424: 3423:Sleep spindle 3421: 3419: 3416: 3414: 3411: 3409: 3406: 3404: 3401: 3399: 3396: 3394: 3391: 3390: 3388: 3386: 3382: 3376: 3373: 3370: 3367: 3365: 3362: 3360: 3357: 3355: 3352: 3350: 3347: 3345: 3342: 3340: 3337: 3334: 3333: 3330: 3327: 3325: 3322: 3320: 3317: 3315: 3312: 3310: 3307: 3305: 3302: 3300: 3297: 3295: 3292: 3290: 3287: 3285: 3282: 3279: 3278: 3276: 3274: 3270: 3264: 3261: 3259: 3256: 3254: 3251: 3249: 3246: 3244: 3241: 3239: 3236: 3235: 3233: 3231:Related tests 3229: 3225: 3218: 3213: 3211: 3206: 3204: 3199: 3198: 3195: 3183: 3180: 3178: 3175: 3173: 3169: 3166: 3164: 3161: 3159: 3156: 3154: 3151: 3149: 3146: 3144: 3141: 3139: 3136: 3134: 3131: 3129: 3126: 3124: 3121: 3119: 3116: 3114: 3111: 3109: 3106: 3104: 3101: 3099: 3096: 3094: 3091: 3087: 3084: 3082: 3079: 3078: 3077: 3074: 3072: 3069: 3067: 3064: 3062: 3059: 3055: 3052: 3050: 3047: 3045: 3042: 3040: 3037: 3035: 3032: 3030: 3027: 3025: 3022: 3021: 3020: 3017: 3016: 3014: 3010: 3000: 2997: 2995: 2992: 2990: 2987: 2985: 2982: 2980: 2977: 2976: 2974: 2970: 2962: 2959: 2958: 2957: 2954: 2950: 2947: 2945: 2942: 2941: 2940: 2937: 2935: 2934:Sleep hygiene 2932: 2930: 2927: 2926: 2924: 2922: 2918: 2912: 2909: 2907: 2904: 2902: 2899: 2897: 2894: 2892: 2891:Sleep inertia 2889: 2887: 2884: 2882: 2879: 2877: 2873: 2870: 2868: 2865: 2863: 2860: 2858: 2855: 2854: 2852: 2848: 2842: 2841:Sleep-talking 2839: 2837: 2836:Sleep driving 2834: 2832: 2829: 2827: 2824: 2822: 2819: 2817: 2814: 2812: 2809: 2808: 2806: 2804: 2800: 2794: 2791: 2789: 2786: 2784: 2781: 2779: 2776: 2774: 2771: 2769: 2766: 2764: 2761: 2760: 2758: 2755: 2748: 2742: 2739: 2737: 2734: 2732: 2729: 2727: 2724: 2722: 2719: 2717: 2714: 2712: 2709: 2707: 2704: 2702: 2699: 2698: 2696: 2694: 2690: 2684: 2681: 2677: 2674: 2672: 2669: 2667: 2664: 2662: 2659: 2657: 2654: 2653: 2652: 2649: 2647: 2644: 2642: 2639: 2638: 2636: 2634: 2630: 2627: 2625: 2621: 2615: 2612: 2610: 2609:Sleep spindle 2607: 2605: 2602: 2600: 2597: 2595: 2592: 2590: 2587: 2585: 2582: 2580: 2577: 2575: 2572: 2570: 2567: 2566: 2564: 2562: 2558: 2552: 2549: 2547: 2544: 2542: 2539: 2538: 2536: 2534: 2529: 2525: 2521: 2514: 2509: 2507: 2502: 2500: 2495: 2494: 2491: 2485: 2482: 2481: 2477: 2470: 2466: 2462: 2458: 2455:(4): 461–84. 2454: 2450: 2445: 2444: 2439: 2435: 2431: 2427: 2423: 2419: 2415: 2411: 2406: 2405: 2400: 2396: 2392: 2388: 2384: 2380: 2377:(7): 923–35. 2376: 2372: 2367: 2366: 2361: 2357: 2353: 2347: 2343: 2339: 2335: 2330: 2329: 2324: 2320: 2316: 2312: 2308: 2304: 2300: 2296: 2291: 2290: 2285: 2281: 2277: 2273: 2269: 2265: 2261: 2257: 2252: 2251: 2246: 2242: 2238: 2234: 2230: 2226: 2222: 2218: 2217:Physiol Behav 2213: 2212: 2206: 2202: 2198: 2194: 2190: 2186: 2180: 2179: 2174: 2170: 2166: 2162: 2158: 2154: 2150: 2146: 2140: 2139: 2134: 2130: 2126: 2122: 2118: 2114: 2110: 2106: 2101: 2100: 2095: 2091: 2086: 2081: 2077: 2073: 2070:(3): 230–53. 2069: 2065: 2061: 2056: 2055: 2050: 2046: 2041: 2036: 2032: 2028: 2024: 2020: 2016: 2011: 2010: 2005: 2001: 1996: 1991: 1987: 1983: 1979: 1975: 1971: 1966: 1965: 1960: 1956: 1952: 1948: 1944: 1940: 1935: 1930: 1927:(7): 936–49. 1926: 1922: 1917: 1913: 1909: 1905: 1901: 1897: 1893: 1889: 1885: 1880: 1879: 1874: 1870: 1866: 1862: 1859:(6): 533–57. 1858: 1854: 1849: 1848: 1843: 1839: 1834: 1829: 1825: 1821: 1817: 1813: 1809: 1804: 1803: 1798: 1794: 1790: 1786: 1782: 1778: 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