According to a popular theory, gamma waves may be implicated in creating the unity of conscious perception (the binding problem). However, there is no agreement on the theory; as researcher C.H. Vanderwolf suggests:
- Whether or not gamma wave activity is related to subjective awareness is a very difficult question which cannot be answered with certainty at the present time.
Gamma waves were initially ignored before the development of digital electroencephalography as analog electroencephalography is restricted to recording and measuring rhythms that are usually less than 25 Hz. One of the earliest reports on them was in 1964 using recordings of the electrical activity of electrodes implanted in the visual cortex of awake monkeys.
Relation to unity of consciousnessEdit
History of ideaEdit
The idea that distinct regions in the brain were being stimulated simultaneously was suggested by the finding in 1988 that two neurons oscillate synchronously (though they are not directly connected) when a single external object stimulates their respective receptive fields. Subsequent experiments by many others demonstrated this phenomenon in a wide range of visual cognition. In particular, Francis Crick and Christof Koch in 1990 argued that there is a significant relation between the binding problem and the problem of visual consciousness and, as a result, that synchronous 40 Hz oscillations may be causally implicated in visual awareness as well as in visual binding. Later the same authors expressed skepticism over the idea that 40 Hz oscillations are a sufficient condition for visual awareness.
A number of experiments conducted by Rodolfo Llinás supports a hypothesis that the basis for consciousness in awake states and dreaming is 40-Hz oscillations throughout the cortical mantle in the form of thalamocortical iterative recurrent activity. In two papers entitled "Coherent 40-Hz oscillation characterizes dream state in humans” (Rodolfo Llinás and Urs Ribary, Proc Natl Acad Sci USA 90:2078-2081, 1993) and "Of dreaming and wakefulness” (Llinas & Pare, 1991), Llinás proposes that the conjunction into a single cognitive event could come about by the concurrent summation of specific and nonspecific 40-Hz activity along the radial dendritic axis of given cortical elements, and that the resonance is modulated by the brainstem and is given content by sensory input in the awake state and intrinsic activity during dreaming. According to Llinás’ hypothesis, known as the thalamocortical dialogue hypothesis for consciousness, the 40-Hz oscillation seen in wakefulness and in dreaming is proposed to be a correlate of cognition, resultant from coherent 40-Hz resonance between thalamocortical-specific and nonspecific loops. In Llinás & Ribary (1993), the authors propose that the specific loops give the content of cognition, and that a nonspecific loop gives the temporal binding required for the unity of cognitive experience.
A lead article by Andreas K. Engel et al. in the journal Consciousness and Cognition (1999) that argues for temporal synchrony as the basis for consciousness, defines the gamma wave hypothesis thus: 
- The hypothesis is that synchronization of neuronal discharges can serve for the integration of distributed neurons into cell assemblies and that this process may underlie the selection of perceptually and behaviorally relevant information.
Role in attentive focusEdit
The suggested mechanism is that gamma waves relate to neural consciousness via the mechanism for conscious attention:
- The proposed answer lies in a wave that, originating in the thalamus, sweeps the brain from front to back, 40 times per second, drawing different neuronal circuits into synch with the precept [sic], and thereby bringing the precept [sic] into the attentional foreground. If the thalamus is damaged even a little bit, this wave stops, conscious awarenesses do not form, and the patient slips into profound coma.
Thus the claim is that when all these neuronal clusters oscillate together during these transient periods of synchronized firing, they help bring up memories and associations from the visual percept to other notions. This brings a distributed matrix of cognitive processes together to generate a coherent, concerted cognitive act, such as perception. This has led to theories that gamma waves are associated with solving the binding problem.
Gamma waves are observed as neural synchrony from visual cues in both conscious and subliminal stimuli. This research also sheds light on how neural synchrony may explain stochastic resonance in the nervous system. Gamma waves are also implicated during rapid eye movement sleep and anesthesia, which involves visualizations.
A 2009 study published in Nature successfully induced gamma waves in mouse brains. Researchers performed this study using optogenetics (the method of combining genetic engineering with light to manipulate the activity of individual nerve cells). The protein channelrhodopsin-2 (ChR2), which sensitizes cells to light, was genetically engineered into these mice, specifically to be expressed in a target-group of interneurons. These fast-spiking (FS) interneurons, known for high electrical activity, were then activated with an optical fiber and laser—the second step in optogenetics. In this way, the cell activity of these interneurons was manipulated in the frequency range of 8–200 Hz. The study produced empirical evidence of gamma wave induction in the approximate interval of 25–100 Hz. The gamma waves were most apparent at a frequency of 40 Hz; this indicates that the gamma waves evoked by FS manipulation are a resonating brain circuit property. This is the first study in which it has been shown that a brain state can be induced through the activation of a specific group of cells. Pushed by the need of understanding how gamma might affect disease pathogenesis, a recent study published in Nature demonstrates that entraining oscillations and spiking at 40 Hz in the hippocampus of a well-established model of Alzheimer's disease (5XFAD mice) reduces Aβ peptides and at the same time activates a microglia response.
Relation to meditationEdit
Experiments on Tibetan Buddhist monks have shown a correlation between transcendental mental states and gamma waves. A suggested explanation is based on the fact that the gamma is intrinsically localized. Neuroscientist Sean O'Nuallain suggests that this very existence of synchronized gamma indicates that something akin to a singularity - or, to be more prosaic, a conscious experience - is occurring. This work adduces experimental and simulated data to show that what meditation masters have in common is the ability to put the brain into a state in which it is maximally sensitive.
As mentioned above, gamma waves have been observed in Tibetan Buddhist monks. A 2004 study took eight long-term Tibetan Buddhist practitioners of meditation and, using electrodes, monitored the patterns of electrical activity produced by their brains as they meditated. The researchers compared the brain activity of the monks to a group of novice meditators (the study had these subjects meditate an hour a day for one week prior to empirical observation). In a normal meditative state, both groups were shown to have similar brain activity. However, when the monks were told to generate an objective feeling of compassion during meditation, their brain activity began to fire in a rhythmic, coherent manner, suggesting neuronal structures were firing in harmony. This was observed at a frequency of 25–40 Hz, the rhythm of gamma waves. These gamma-band oscillations in the monk’s brain signals were the largest seen in humans (apart from those in states such as seizures). Conversely, these gamma-band oscillations were scant in novice meditators. Though, a number of rhythmic signals did appear to strengthen in beginner meditators with further experience in the exercise, implying that the aptitude for one to produce gamma-band rhythm is trainable.
Such evidence and research in gamma-band oscillations may explain the heightened sense of consciousness, bliss, and intellectual acuity subsequent to meditation. Notably, meditation is known to have a number of health benefits: stress reduction, mood elevation, and increased life expectancy of the mind and its cognitive functions. The current Dalai Lama meditates for four hours each morning, and he says that it is hard work. He elaborates that if neuroscience can construct a way in which he can reap the psychological and biological rewards of meditation without going through the practice each morning, he would be apt to adopt the innovation.
Many neuroscientists are not convinced of the gamma wave argument. Arguments against it range from the possibility of mismeasurement – it has been suggested that EEG-measured gamma waves could be in many cases an artifact of electromyographic activity – to relations to other neural function, such as minute eye movements.
Moreover, recent studies using magnetoencephalography (MEG), which does not suffer the potential artifacts associated with EEG, have identified gamma activity associated with sensory processing, mainly in the visual cortex.
A recent study with micro-electrodes in monkey and human showed that gamma oscillations are present and are clearly correlated with the firing of single neurons, mostly inhibitory neurons. The gamma oscillations were found in humans during all states of the wake-sleep cycle, and were maximally coherent during slow-wave sleep.
Bearing this theory in mind, a number of questions remain unexplained regarding details of exactly how the temporal synchrony results in a conscious awareness or how a new percept "calls for" the synchrony, etc.
- Hughes JR (July 2008). "Gamma, fast, and ultrafast waves of the brain: their relationships with epilepsy and behavior". Epilepsy Behav. 13 (1): 25–31. doi:10.1016/j.yebeh.2008.01.011. PMID 18439878.
- Ian Gold (1999). "Does 40-Hz oscillation play a role in visual consciousness?". Consciousness and Cognition. 8 (2): 186–195. doi:10.1006/ccog.1999.0399. PMID 10448001.
- Buzsaki, György (2006). "Cycle 9, The Gamma Buzz". Rhythms of the brain. Oxford. ASIN 0195301064.
- Robert Pollack, The Missing Moment, 1999
- Singer, W.; Gray, C.M. (1995). "Visual feature integration and the temporal correlation hypothesis". Annu. Rev. Neurosci. 18: 555–586. doi:10.1146/annurev.ne.18.030195.003011. PMID 7605074.
- Vanderwolf CH (Feb 2000). "Are neocortical gamma waves related to consciousness?". Brain Res. 855 (2): 217–24. doi:10.1016/S0006-8993(99)02351-3. PMID 10677593.
- Hughes JR. (1964). Responses from the visual cortex of unanesthetized monkeys. pp. 99–153. In: Pfeiffer CC, Smythies JR, (Eds), International review of neurobiology vol. 7, Academic Press, New York OCLC 43986646
- Crick, F., & Koch, C. (1990b). Towards a neurobiological theory of consciousness. Seminars in the Neurosciences v.2, 263-275.
- Crick, F., Koch, C. (2003). "Framework for consciousness". Nature Neuroscience. 6 (2): 119–26. doi:10.1038/nn0203-119. PMID 12555104.
- Andreas K. Engel; Pascal Fries; Peter Koenig; Michael Brecht; Wolf Singer (1999). "Temporal Binding, Binocular Rivalry, and Consciousness". Consciousness and Cognition. 8 (2): 128–151. doi:10.1006/ccog.1999.0389.
- Melloni L, Molina C, Pena M, Torres D, Singer W, Rodriguez E (Mar 2007). "Synchronization of neural activity across cortical areas correlates with conscious perception". J Neurosci. 27 (11): 2858–65. doi:10.1523/JNEUROSCI.4623-06.2007. PMID 17360907.
- Siegel M, Donner TH, Oostenveld R, Fries P, Engel AK (Mar 2008). "Neuronal synchronization along the dorsal visual pathway reflects the focus of spatial attention". Neuron. 60 (4): 709–719. doi:10.1016/j.neuron.2008.09.010. PMID 19038226.
- Gregoriou GG, Gotts SJ, Zhou H, Desimone R (Mar 2009). "High-frequency, long-range coupling between prefrontal and visual cortex during attention". Science. 324 (5931): 1207–1210. Bibcode:2009Sci...324.1207G. doi:10.1126/science.1171402. PMC . PMID 19478185.
- Baldauf D, Desimone R (Mar 2014). "Neural mechanisms of object-based attention". Science. 344 (6182): 424–427. Bibcode:2014Sci...344..424B. doi:10.1126/science.1247003. PMID 24763592.
- Ward LM, Doesburg SM, Kitajo K, MacLean SE, Roggeveen AB (Dec 2006). "Neural synchrony in stochastic resonance, attention, and consciousness". Can J Exp Psychol. 60 (4): 319–26. doi:10.1037/cjep2006029. PMID 17285879.
- <J. Cardin, M. Carle, K. Meletis, U. Knoblich, F. Zhang, K. Deisseroth, Li-Huei Tsai and Christopher Moore (2009) Driving fast-spiking cells induces gamma rhythm and controls sensory responses. Nature, 459: 663-668.>
- Iaccarino, Hannah F.; Singer, Annabelle C.; Martorell, Anthony J.; Rudenko, Andrii; Gao, Fan; Gillingham, Tyler Z.; Mathys, Hansruedi; Seo, Jinsoo; Kritskiy, Oleg; Abdurrob, Fatema; Adaikkan, Chinnakkaruppan; Canter, Rebecca G.; Rueda, Richard; Brown, Emery N.; Boyden, Edward S.; Tsai, Li-Huei (7 December 2016). "Gamma frequency entrainment attenuates amyloid load and modifies microglia". Nature. 540 (7632): 230–235. Bibcode:2016Natur.540..230I. doi:10.1038/nature20587.
- O'Nuallain, Sean. "Zero Power and Selflessness: What Meditation and Conscious Perception Have in Common". Retrieved 2009-05-30. Journal: Cognitive Sciences 4(2).
- Kaufman, Marc (January 3, 2005). "Meditation Gives Brain a Charge, Study Finds". The Washington Post. Retrieved May 3, 2010.
- Lutz A.; Greischar L.L.; Rawlings N.B.; Ricard M.; Davidson R.J. (2004). "Long-term meditators self-induce high amplitude gamma synchrony during mental practice". Proceedings of the National Academy of Sciences USA. 101: 16369–16373. Bibcode:2004PNAS..10116369L. doi:10.1073/pnas.0407401101. PMC . PMID 15534199.
- "Scientific American:Meditation On Demand".
- Whitham EM, Pope KJ, Fitzgibbon SP, et al. (Aug 2007). "Scalp electrical recording during paralysis: quantitative evidence that EEG frequencies above 20 Hz are contaminated by EMG". Clin Neurophysiol. 118 (8): 1877–88. doi:10.1016/j.clinph.2007.04.027. PMID 17574912.
- Whitham EM, Lewis T, Pope KJ, et al. (May 2008). "Thinking activates EMG in scalp electrical recordings". Clin Neurophysiol. 119 (5): 1166–75. doi:10.1016/j.clinph.2008.01.024. PMID 18329954.
- Yuval-Greenberg S, Tomer O, Keren AS, Nelken I, Deouell LY (May 2008). "Transient induced gamma-band response in EEG as a manifestation of miniature saccades". Neuron. 58 (3): 429–41. doi:10.1016/j.neuron.2008.03.027. PMID 18466752.
- Dynamic predictions: Oscillations and synchrony in top-down processing, AK Engel, P Fries, W Singer, Nature Reviews Neuroscience, 2001
- Kort, N; Cuesta, P; Houde, JF; Nagarajan, SS (2016). "Bihemispheric network dynamics coordinating vocal feedback control". Human Brain Mapping. 37 (4): 1474–1485. doi:10.1002/hbm.23114.
- Adjamian, P; Holliday, IE; Barnes, GR; Hillebrand, A; Hadjipapas, A; Singh, KD (2004). "Induced stimulus-dependent Gamma oscillations in visual stress". European Journal of Neuroscience. 20: 587–592. doi:10.1111/j.1460-9568.2004.03495.x.
- Hadjipapas A.; Adjamian P; Swettenham J.B.; Holliday I.E.; Barnes G.R. (2007). "Stimuli of varying spatial scale induce gamma activity with distinct temporal characteristics in human visual cortex". NeuroImage. 35 (2): 518–30. doi:10.1016/j.neuroimage.2007.01.002.
- Muthukumaraswamy SD, Singh KD (2008). "Spatiotemporal frequency tuning of BOLD and gamma band MEG responses compared in primary visual cortex". NeuroImage. 40: 1552–1560. doi:10.1016/j.neuroimage.2008.01.052. PMID 18337125.
- Swettenham JB, Muthukumaraswamy SD, Singh KD (2009). "Spectral properties of induced and evoked gamma oscillations in human early visual cortex to moving and stationary stimuli". Journal of Neurophysiology. 102: 1241–1253. doi:10.1152/jn.91044.2008. PMID 19515947.
- Le Van Quyen M.; Muller L.E.; Telenczuk B.; Halgren E.; Cash S.; Hatsopoulos N.; Dehghani N.; Destexhe A. (2016). "High-frequency oscillations in human and monkey neocortex during the wake-sleep cycle". Proceedings of the National Academy of Sciences USA. 113 (33): 93639368. doi:10.1073/pnas.1523583113. PMC . PMID 27482084.