Neuromodulation of brain state and behavior
Open accessAbstract
Neural activity and behavior are both notoriously variable, with responses differing widely between repeated presentation of identical stimuli or trials. Recent results in humans and animals reveal that these variations are not random in their nature, but may in fact be due in large part to rapid shifts in neural, cognitive, and behavioral states. Here we review recent advances in the understanding of rapid variations in the waking state, how variations are generated, and how they modulate neural and behavioral responses in both mice and humans. We propose that the brain has an identifiable set of states through which it wanders continuously in a nonrandom fashion, owing to the activity of both ascending modulatory and fast-acting corticocortical and subcortical-cortical neural pathways. These state variations provide the backdrop upon which the brain operates, and understanding them is critical to making progress in revealing the neural mechanisms underlying cognition and behavior.
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Sleep is a brain state characterized by large-scale organization of neuronal activity capturing virtually all brain regions.
“Sleep is a brain state that is characterized by a large-scale organization of neuronal activity that captures virtually all brain regions”
From Sleep—A brain-state serving systems memory consolidation (2023) · cites this paper as [8]
The Introduction (page unavailable) describes brain states in terms of neuronal and regional activity, and Cellular and Network Mechanisms of State-Dependent Activity says slow oscillations are prevalent during sleep. But the latter describes oscillations generated in local cortical regions; our copy does not establish that sleep organizes neuronal activity across virtually all brain regions. (LLM, conf=0.76)
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Sleep—A brain-state serving systems memory consolidation
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Natural behavior is the language of the brain
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Noradrenergic modulation of rhythmic neural activity shapes selective attention
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“Hence, acetylcholine promotes cortical desynchronization, similar to the action of noradrenaline [9,11,31,83].”
BackgroundInfluentialWhy is everyone talking about brain state?
Greene, A. S., Horien, C. L., Barson, D. G., Scheinost, D., & Constable, R. T. (2023) · Trends in Neurosciences · doi:10.1016/j.tins.2023.04.001 · Cited by 148 · see citing works
“Further, increased network integration is a hallmark of increased arousal, as indexed by pupil size [39], consistent with the idea that increased arousal and related brain activity track increased attention and task performance [48], although only to a point [4] (next section).”
BackgroundVagus nerve stimulation induces widespread cortical and behavioral activation
Collins, L. N., Boddington, L. J., Steffan, P. J., & McCormick, D. A. (2021) · Current Biology · doi:10.1016/j.cub.2021.02.049 · Cited by 147 · see citing works
Self-citationAstrocyte regulation of neural circuit activity and network states
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“current understanding of the mechanisms underlying brain states usually disregards the participation of astrocytes (e.g., Lee & Dan, 2012; McCormick et al., 2020; Zagha & McCormick, 2014), which, however,”
BackgroundInfluentialPhasic Activation of Dorsal Raphe Serotonergic Neurons Increases Pupil Size
Cazettes, F., Reato, D., Morais, J. P., Renart, A., & Mainen, Z. F. (2020) · Current Biology · doi:10.1016/j.cub.2020.09.090 · Cited by 133 · see citing works
Brain-inspired learning in artificial neural networks: A review
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Pupil diameter is not an accurate real-time readout of locus coeruleus activity
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“Introduction Fluctuations of brain states, such as arousal and attention, strongly impact sensory processing, decision- making, and animal behavior (Harris and Thiele, 2011; Lee and Dan, 2012; Thiele and Bellgrove, 2018; Petersen, 2019; McCormick et al., 2020).”
BackgroundGrimm, C., Duss, S. N., Privitera, M., Munn, B., Karalis, N., Frässle, S., Wilhelm, M., Patriarchi, T., Razansky, D., Wenderoth, N., et al. (2024) · Nature Neuroscience · doi:10.1038/s41593-024-01755-8 · Cited by 128 · see citing works
“Spatially coordinated, temporally sustained patterns of neural activity give rise to brain activity patterns that can change discretely or continuously, with or without external stimuli 45 .”
BackgroundVisual thalamocortical mechanisms of waking state-dependent activity and alpha oscillations
Nestvogel, D., & McCormick, D. A. (2021) · Neuron · doi:10.1016/j.neuron.2021.10.005 · Cited by 95 · see citing works
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Self-citationComplexity and 1/f slope jointly reflect brain states
Medel, V., Irani, M., Crossley, N. A., Ossandón, T., & Boncompte, G. (2023) · Scientific Reports · doi:10.1038/s41598-023-47316-0 · Cited by 85 · see citing works
Deco, G., Perl, Y. S., Bocaccio, H., Tagliazucchi, E., & Kringelbach, M. L. (2022) · Communications Biology · doi:10.1038/s42003-022-03505-7 · Cited by 81 · see citing works
“H24-2-203(4)) approved all experimental and surgical procedures and the research carefully followed the recommendations of the Weatherall report “The use of non-human primates in research”.”
BackgroundMethodInfluentialOffline memory consolidation during waking rest
Wamsley, E. J. (2022) · Nature Reviews Psychology · doi:10.1038/s44159-022-00072-w · Cited by 76 · see citing works
Centripetal integration of past events in hippocampal astrocytes regulated by locus coeruleus
Rupprecht, P., Duss, S. N., Becker, D., Lewis, C., Bohacek, J., & Helmchen, F. (2024) · Nature Neuroscience · doi:10.1038/s41593-024-01612-8 · Cited by 79 · see citing works
“Our study suggests that the disen-tanglement of these processes, which has turned out to be challenging for neurons 34 , is not easier for astrocytes.”
Arousal as a universal embedding for spatiotemporal brain dynamics
Raut, R. V., Rosenthal, Z. P., Wang, X., Miao, H., Zhang, Z., Lee, J., Raichle, M. E., Bauer, A. Q., Brunton, S. L., Brunton, B. W., et al. (2025) · Nature · doi:10.1038/s41586-025-09544-4 · Cited by 79 · see citing works
“The first, primarily involving invasive recordings in awake mice, has shown that cortical activity (especially above 1 Hz) is modulated in a region-dependent manner according to ongoing fluctuations in arousal and behavioural state 1,3,12 ; however, general principles have remained elusive 12,15 .”
InfluentialSpectral signature and behavioral consequence of spontaneous shifts of pupil-linked arousal in human
Podvalny, E., King, L. E., & He, B. J. (2021) · eLife · doi:10.7554/elife.68265 · Cited by 66 · see citing works
“68265 2 of 25 and event- related pupillary response (Goldwater, 1972) that are likely governed by distinct neuromodulatory circuits and/or modes of brain activity (McCormick et al., 2020; Joshi and Gold, 2020; AstonJones and Cohen, 2005).”
BackgroundInfluentialSelf-citationTowards a pluralistic neurobiological understanding of consciousness
He, B. J. (2023) · Trends in Cognitive Sciences · doi:10.1016/j.tics.2023.02.001 · Cited by 73 · see citing works
Self-citationDistinct ensembles in the noradrenergic locus coeruleus are associated with diverse cortical states
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Unifying turbulent dynamics framework distinguishes different brain states
Escrichs, A., Perl, Y. S., Uribe, C., Càmara, E., Türker, B., Pyatigorskaya, N., López‐González, A., Pallavicini, C., Panda, R., Annen, J., et al. (2022) · Communications Biology · doi:10.1038/s42003-022-03576-6 · Cited by 65 · see citing works
“…remain unsolved, and most importantly, there is a need to arrive at an agreed definition of brain states (Carhart-Harris et al., 2016; Deco et al., 2015; Gervasoni et al., 2004; Kringelbach & Deco, 2020; McCormick et al., 2020; Northoff, 2013; Tagliazucchi et al., 2016; G Tononi et al., 1994).”
BackgroundCognition from the Body-Brain Partnership: Exaptation of Memory
Buzsáki, G., & Tingley, D. (2023) · Annual Review of Neuroscience · doi:10.1146/annurev-neuro-101222-110632 · Cited by 62 · see citing works
“Switching between these states is triggered or at least correlated with high and low release of subcortical neuromodulators, respectively (McCormick et al. 2020).”
BackgroundOlfactory modulation of the medial prefrontal cortex circuitry: Implications for social cognition
Bhattarai, J. P., Etyemez, S., Jaaro-Peled, H., Janke, E., Tolosa, U. D. L., Kamiya, A., Gottfried, J. A., Sawa, A., & Ma, M. (2021) · Seminars in Cell and Developmental Biology · doi:10.1016/j.semcdb.2021.03.022 · Cited by 56 · see citing works
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“One of the factors that contributes to this behavioral variability is the arousal state of the brain, which fluctuates spontaneously over time, causing variability in neuronal responsiveness to sensory stimulation (also often referred to as brain state) (Greene et al., 2023; McCormick et al., 2020; Waschke et al., 202…”
Sleep microstructure organizes memory replay
Chang, H., Tang, W., Wulf, A. M., Nyasulu, T., Wolf, M. E., Fernández‐Ruiz, A., & Oliva, A. (2025) · Nature · doi:10.1038/s41586-024-08340-w · Cited by 64 · see citing works
“In both humans and animals, pupil fluctuations have been proven as a sensitive marker of brain-state changes 23 , and thus can be a useful readout of sleep dynamics; yet, data about pupil dynamics in rodents during NREM sleep are scarce.”
Yang, T., Shen, T., Duan, B., Liu, Z., & Wang, C. (2025) · ACS Sensors · doi:10.1021/acssensors.4c03314 · Cited by 58 · see citing works
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“These states can be physiological (e.g., awake, slow wave sleep, REM), drug-induced (e.g., anesthesia), or even different attentional states or variations within the awake state (McCormick et al., 2020).”
BackgroundTheriault, J. E., Shaffer, C., Dienel, G. A., Sander, C. Y. M., Hooker, J. M., Dickerson, B. C., Barrett, L. F., & Quigley, K. S. (2023) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2023.105373 · Cited by 55 · see citing works
Decision-making dynamics are predicted by arousal and uninstructed movements
Hulsey, D., Zumwalt, K., Mazzucato, L., McCormick, D. A., & Jaramillo, S. (2024) · Cell Reports · doi:10.1016/j.celrep.2024.113709 · Cited by 54 · see citing works
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Multimodal EEG-fMRI: advancing insight into large-scale human brain dynamics
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Innovation at the Intersection: Emerging Translational Research in Neurology and Psychiatry
Tanaka, M., Battaglia, S., Giménez‐Llort, L., Chen, C., Hepsomali, P., Avenanti, A., & Vécsei, L. (2024) · Cells · doi:10.3390/cells13100790 · Cited by 48 · see citing works
“Neuromodulation is the process of altering neural activity and function by applying external stimuli such as light, electric currents, magnetic fields, or drugs to specific brain regions or networks [14,191].”
BackgroundArousal impacts distributed hubs modulating the integration of brain functional connectivity
Lee, K., Horien, C. L., O’Connor, D. A., Garand-Sheridan, B., Tokoglu, F., Scheinost, D., Lake, E. M. R., & Constable, R. T. (2022) · NeuroImage · doi:10.1016/j.neuroimage.2022.119364 · Cited by 38 · see citing works
“changes in pupil size, without a need to place EEG electrodes on the scalp (Joshi and Gold, 2020; Liu and Falahpour, 2020; McCormick et al., 2020).”
Active Dendrites and Local Field Potentials: Biophysical Mechanisms and Computational Explorations
Sinha, M., & Narayanan, R. (2021) · Neuroscience · doi:10.1016/j.neuroscience.2021.08.035 · Cited by 42 · see citing works
“Importantly, independent MPMOSS or GA simulations could be per-1392 formed to generate distinct populations of models for stimuli (Getting, 1989; Marder and Thirumalai, 2002; Lee and Dan, 2012; Marder, 2012; Bargmann and Marder, 2013; Marder et al., 2014; McCormick et al., 2020).”
van den Berg, M., Toen, D. V., Verhoye, M., & Keliris, G. A. (2023) · Frontiers in Aging Neuroscience · doi:10.3389/fnagi.2023.1081058 · Cited by 41 · see citing works
The predictive nature of spontaneous brain activity across scales and species
Dimakou, A., Pezzulo, G., Zangrossi, A., & Corbetta, M. (2025) · Neuron · doi:10.1016/j.neuron.2025.02.009 · Cited by 43 · see citing works
Bioelectronic Medicine: a multidisciplinary roadmap from biophysics to precision therapies
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“It should be noted that in addition, neuromodulators released at synapses further shape synaptic transmission (O’Brien, 2014, 2019; Alcedo and Prahlad, 2020; McCormick et al., 2020; Wu et al., 2022).”
BackgroundSharing neurophysiology data from the Allen Brain Observatory
de Vries, S. E. J., Siegle, J. H., & Koch, C. (2023) · eLife · doi:10.7554/elife.85550 · Cited by 43 · see citing works
“Whereas gene expression and connectivity are expected to change relatively slowly, neural responses in awake subjects can vary dramatically from moment to moment, even during apparently quiescent periods ( McCormick et al., 2020 ).”
BackgroundA common thalamic hub for general and defensive arousal control
Wang, Y., You, L., Tan, K., Li, M., Zou, J., Zhao, Z., Hu, W., Li, T., Xie, F., Li, C., et al. (2023) · Neuron · doi:10.1016/j.neuron.2023.07.007 · Cited by 38 · see citing works
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“1-4 High-magnification images of areas indicated by dashed boxes showing the cortex (1), the hippocampus (2), the BLA (3), and the hypothalamus (4).”
BackgroundThe lack of temporal brain dynamics asymmetry as a signature of impaired consciousness states
Guzman, E. G., Perl, Y. S., Vohryzek, J., Escrichs, A., Manasova, D., Türker, B., Tagliazucchi, E., Kringelbach, M. L., Sitt, J. D., & Deco, G. (2023) · Interface Focus · doi:10.1098/rsfs.2022.0086 · Cited by 34 · see citing works
Neurobiological Underpinnings of Hyperarousal in Depression: A Comprehensive Review
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“This may partly be due to incomprehensive investigations regarding the neural basis of arousal—key features of the brain states—in depression [1].”
BackgroundState-related neural influences on fMRI connectivity estimation
Martin, C. G., He, B. J., & Chang, C. E. (2021) · NeuroImage · doi:10.1016/j.neuroimage.2021.118590 · Cited by 33 · see citing works
“…l f y t l p 2 2 v F d n d a r 2 q c s a g a n m s 2 B c s 2 t t T m s L r n f r c S ( c i r al conditions and vary as a function of physiological or psychological tate, either externally or internally driven ( Thayer and Lane, 2000 ; ritchley, 2005 ; McCraty and Zayas, 2014 ; Chand et al., 2020 ).”
BackgroundSelf-citationShi, Y.-L., Steinmetz, N. A., Moore, T., Boahen, K. A., & Engel, T. A. (2022) · Nature Communications · doi:10.1038/s41467-021-27724-4 · Cited by 31 · see citing works
Alpha EEG Activity and Pupil Diameter Coupling during Inactive Wakefulness in Humans
Montefusco‐Siegmund, R., Schwalm, M., Jubal, E. R., Devia, C., Egaña, J. I., & Maldonado, P. (2022) · eNeuro · doi:10.1523/eneuro.0060-21.2022 · Cited by 33 · see citing works
“A parsimonious explanation postulates that the internal system is continuously changing, and so, however similar the external environment may be, those internal fluctuations affect the individual differently at each moment (Lee and Dan, 2012; McCormick et al., 2020).”
BackgroundComplementary roles of serotonergic and cholinergic systems in decisions about when to act
Khalighinejad, N., Manohar, S., Husain, M., & Rushworth, M. F. S. (2022) · Current Biology · doi:10.1016/j.cub.2022.01.042 · Cited by 29 · see citing works
“This is supported by emerging evidence that ACh can rapidly and selectively modulate activity in specific brain areas.(31,33,35) Here, we showed that increasing systemic ACh invigorated movements so that animals acted faster when offered a reward as compared with the control condition.”
BackgroundThe vertical light-gradient and its potential impact on animal distribution and behavior
Nilsson, D., Smolka, J., & Bok, M. J. (2022) · Frontiers in Ecology and Evolution · doi:10.3389/fevo.2022.951328 · Cited by 32 · see citing works
“” This concept is gaining ground in both ecology and neurobiology (Berman et al., 2016; Gurarie et al., 2016; Mahoney and Young, 2017; Naässel et al., 2019; Russell et al., 2019; McCormick et al., 2020).”
BackgroundSörensen, L. K. A., Bohté, S. M., Slagter, H. A., & Scholte, H. S. (2022) · PLoS Computational Biology · doi:10.1371/journal.pcbi.1009976 · Cited by 29 · see citing works
“These increases in gain were suggested to be a result of a global release of noradrenaline (in conjunction with other neuromodulators [48]), making them spatially unspecific [4,14,16,44,48].”
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Stable continual learning through structured multiscale plasticity manifolds
Mishra, P., & Narayanan, R. (2021) · Current Opinion in Neurobiology · doi:10.1016/j.conb.2021.07.009 · Cited by 31 · see citing works
Functional Coupling of the Locus Coeruleus Is Linked to Successful Cognitive Control
Grueschow, M., Kleim, B., & Ruff, C. C. (2022) · Brain Sciences · doi:10.3390/brainsci12030305 · Cited by 30 · see citing works
“inform human studies to test the functional and behavioral relevance of the neural arousal circuits [42].”
BackgroundMorabito, A., Zerlaut, Y., Serraz, B., Sala, R. W., Paoletti, P., & Rebola, N. (2022) · Cell Reports · doi:10.1016/j.celrep.2022.110415 · Cited by 30 · see citing works
Network mechanisms of ongoing brain activity’s influence on conscious visual perception
Wu, Y., Podvalny, E., Levinson, M., & He, B. J. (2024) · Nature Communications · doi:10.1038/s41467-024-50102-9 · Cited by 30 · see citing works
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“…network reorganization in the context of an auditory oddball task, we speculate that the CS-INR system might play a more general role in cognitive functions, such as adaptations in environmental volatility [53, 76], brain state switches/variations [44, 77], cognitive control [15, 37, 48, 78].”
BackgroundThe influence of cortical activity on perception depends on behavioral state and sensory context
Russell, L. E., Fişek, M., Yang, Z., Tan, L. P., Packer, A. M., Dalgleish, H. W. P., Chettih, S. N., Harvey, C. D., & Häusser, M. (2024) · Nature Communications · doi:10.1038/s41467-024-46484-5 · Cited by 54 · see citing works
“The perception of a sensory stimulus is modulated by the behavioral state in whichit isexperienced – successfuldetection of a stimulus can be increased during periods of arousal or attention 1,2 .”
BackgroundImaging the temporal dynamics of brain states with highly sampled fMRI
Yang, Z., & Lewis, L. D. (2021) · Current Opinion in Behavioral Sciences · doi:10.1016/j.cobeha.2021.02.005 · Cited by 28 · see citing works
Control of Brain State Transitions with a Photoswitchable Muscarinic Agonist
Barbero‐Castillo, A., Riefolo, F., Matera, C., Caldas‐Martínez, S., Mateos‐Aparicio, P., Weinert, J. F., Garrido‐Charles, A., Claro, E., Sanchez-Vives, M. V., & Gorostiza, P. (2021) · Advanced Science · doi:10.1002/advs.202005027 · Cited by 28 · see citing works
Chang, Y., Yep, R., & Wang, C. (2024) · Psychophysiology · doi:10.1111/psyp.14726 · Cited by 27 · see citing works
“…on a trial- by-trial basis, is associated with fluctuations in various neural activity and brain states (Joshi et al., 2016; Joshi & Gold, 2020; McCormick et al., 2020; McGinley et al., 2015; Reimer et al., 2014, 2016; Vinck et al., 2015), and is increasingly adopted as an indicator of…”
InfluentialSleep’s contribution to memory formation
Lutz, N. D., Harkotte, M., & Jan Born, A. (2025) · Physiological Reviews · doi:10.1152/physrev.00054.2024 · Cited by 40 · see citing works
A Causal Network Analysis of Neuromodulation in the Mood Processing Network
Qiao, S., Sedillo, J. I., Brown, K. A., Ferrentino, B., & Pesaran, B. (2020) · Neuron · doi:10.1016/j.neuron.2020.06.012 · Cited by 27 · see citing works
“By decoding the specific state of modulator networks associated with disordered multiregional communication (Braun et al., 2018; McCormick et al., 2020; Shanechi, 2019), our results offer a new strategy for restoring disordered multiregional communication in a novel BMI that decodes the state of modulator networks to…”
BackgroundChanging the Cortical Conductor’s Tempo: Neuromodulation of the Claustrum
Wong, K. L. L., Nair, A., & Augustine, G. J. (2021) · Frontiers in Neural Circuits · doi:10.3389/fncir.2021.658228 · Cited by 26 · see citing works
“They are thought to result from the actions of neuromodulators, such as acetylcholine (ACh), dopamine (DA) and serotonin (5-HT), that affect brain processes ranging from macroscopic networks down to subcellular signaling (Kringelbach and Deco, 2020; McCormick et al., 2020).”
BackgroundSobczak, F., Pais‐Roldán, P., Takahashi, K., & Yu, X. (2021) · eLife · doi:10.7554/elife.68980 · Cited by 22 · see citing works
Neuromodulation and Individuality
Maloney, R. T. (2021) · Frontiers in Behavioral Neuroscience · doi:10.3389/fnbeh.2021.777873 · Cited by 23 · see citing works
“Neuromodulators have strong effects on behaviors, and within an animal shifts in neuromodulators are a driver of trial to trial variability (McCormick et al., 2020).”
BackgroundA dynamical systems view of neuroethology: Uncovering stateful computation in natural behaviors
Robson, D. N., & Li, J. (2022) · Current Opinion in Neurobiology · doi:10.1016/j.conb.2022.01.002 · Cited by 26 · see citing works
Deciphering neuronal variability across states reveals dynamic sensory encoding
Akella, S., Ledochowitsch, P., Siegle, J. H., Belski, H., Denman, D. J., Buice, M. A., Durand, S., Koch, C., Olsen, S. R., & Jia, X. (2025) · Nature Communications · doi:10.1038/s41467-025-56733-w · Cited by 23 · see citing works
“Brain state variations often exhibit strong correlations with the animal’s behavioral context (Mccormick et al. 2020; Zagha and McCormick 2014).”
InfluentialWalking humans and running mice: perception and neural encoding of optic flow during self-motion
Horrocks, E. A. B., Mareschal, I., & Saleem, A. B. (2022) · Philosophical Transactions of the Royal Society B Biological Sciences · doi:10.1098/rstb.2021.0450 · Cited by 25 · see citing works
Memory Consolidation during Ultra-short Offline States
Wamsley, E. J., Arora, M., Gibson, H., Powell, P., & Collins, M. B. (2023) · Journal of Cognitive Neuroscience · doi:10.1162/jocn_a_02035 · Cited by 22 · see citing works
Time, space, memory and brain–body rhythms
Buzsáki, G. (2025) · Nature reviews. Neuroscience · doi:10.1038/s41583-025-00987-2 · Cited by 28 · see citing works
“In humans, an ~1 min rhythm generated by these cells phase-modulates the power of the cortical alpha rhythm in the right anterior interoceptive insular cortex 158 .”
Goc, G. L., Lafaye, J., Karpenko, S., Bormuth, V., Candelier, R., & Debrégeas, G. (2021) · BMC Biology · doi:10.1186/s12915-021-01126-w · Cited by 18 · see citing works
“patterns of persistent neural activity [5, 6].”
The Neuromodulatory Basis of Aggression: Lessons From the Humble Fruit Fly
Palavicino‐Maggio, C. B., & Sengupta, S. (2022) · Frontiers in Behavioral Neuroscience · doi:10.3389/fnbeh.2022.836666 · Cited by 18 · see citing works
Spiking activity in the visual thalamus is coupled to pupil dynamics across temporal scales
Crombie, D., Spacek, M. A., Leibold, C., & Busse, L. (2024) · PLoS Biology · doi:10.1371/journal.pbio.3002614 · Cited by 20 · see citing works
“More generally, a substantial fraction of shared variability present in large populations of neurons in visual cortex can be explained by high-dimensional sets of behaviors [19], indicating that the state of sensory systems at any given moment results from a combination of multiple processes [3].”
BackgroundLin, Z.-J., Gu, X., Gong, W., Wang, M., Wu, Y., Wang, Q., Wu, X., Zhao, X.-Y., Zhu, M. X., Wang, L., et al. (2024) · Journal of Clinical Investigation · doi:10.1172/jci181095 · Cited by 19 · see citing works
Look-up and look-down neurons in the mouse visual thalamus during freely moving exploration
Orlowska‐Feuer, P., Ebrahimi, A. S., Zippo, A. G., Petersen, R. S., Lucas, R. J., & Storchi, R. (2022) · Current Biology · doi:10.1016/j.cub.2022.07.049 · Cited by 19 · see citing works
In Situ Synthesis and Assembly of Functional Materials and Devices in Living Systems
Wang, W., Sessler, C. D., Wang, X., & Jia Liu, B. (2024) · Accounts of Chemical Research · doi:10.1021/acs.accounts.4c00049 · Cited by 18 · see citing works
Links between the brain and body during sleep: implications for memory processing
Whitehurst, L. N., Subramoniam, A., Krystal, A. D., & Prather, A. A. (2022) · Trends in Neurosciences · doi:10.1016/j.tins.2021.12.007 · Cited by 18 · see citing works
“Now, feedback networks between the hippocampus and neocortex become critical for replay and transfer [84 – 87].”
BackgroundBrain Infraslow Activity Correlates With Arousal Levels
Sihn, D., & Kim, S.-P. (2022) · Frontiers in Neuroscience · doi:10.3389/fnins.2022.765585 · Cited by 18 · see citing works
Andrews, L., Keller, S. S., Osman‐Farah, J., & Macerollo, A. (2023) · Brain Communications · doi:10.1093/braincomms/fcad171 · Cited by 17 · see citing works
“Given that neuromodulatory effects have been shown to be sensitive to changes in brain state, 122,123 it would make sense that the structural architecture of the brain—in terms of grey and white matter integrity —would instantiate a mediating factor for therapeutic efficacy.”
Locus Coeruleus firing patterns selectively modulate brain activity and dynamicsPreprint
Grimm, C., Duss, S. N., Privitera, M., Munn, B., Frässle, S., Chernysheva, M., Patriarchi, T., Razansky, D., Wenderoth, N., Shine, J. M., et al. (2022) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2022.08.29.505672 · Cited by 18 · see citing works
Acetylcholine engages distinct amygdala microcircuits to gate internal theta rhythm
Bratsch-Prince, J. X., Warren, J., Jones, G. C., McDonald, A. J., & Mott, D. D. (2024) · Journal of Neuroscience · doi:10.1523/jneurosci.1568-23.2024 · Cited by 16 · see citing works
A thalamocortical pathway controlling impulsive behavior
Guzulaitis, R., & Palmer, L. M. (2023) · Trends in Neurosciences · doi:10.1016/j.tins.2023.09.001 · Cited by 16 · see citing works
A time window for memory consolidation during NREM sleep revealed by cAMP oscillation
Deng, Z., Fei, X., Zhang, S., & Xu, M. (2025) · Neuron · doi:10.1016/j.neuron.2025.03.020 · Cited by 17 · see citing works
Neural circuit underlying individual differences in visual escape habituation
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Averna, A., Marceglia, S. R. F., Priori, A., & Foffani, G. (2022) · npj Parkinson s Disease · doi:10.1038/s41531-022-00399-4 · Cited by 16 · see citing works
“(1) and (2) respectively) by varying either the amplitude or the frequency of a sinusoidal carrier signal fc with a modulating signal of lower frequency fm (Lathi and Ding, 2010):”
BackgroundNeurobiological reduction: From cellular explanations of behavior to interventions
Parker, D. (2022) · Frontiers in Psychology · doi:10.3389/fpsyg.2022.987101 · Cited by 18 · see citing works
Toward a computational role for locus coeruleus/norepinephrine arousal systems
Nassar, M. R. (2024) · Current Opinion in Behavioral Sciences · doi:10.1016/j.cobeha.2024.101407 · Cited by 15 · see citing works
Papadopoulos, L., Jo, S., Zumwalt, K., Wehr, M., Jaramillo, S., McCormick, D. A., & Mazzucato, L. (2025) · Neuron · doi:10.1016/j.neuron.2025.11.011 · Cited by 15 · see citing works
Self-citationDissociable Components of Information Encoding in Human Perception
Vidaurre, D., Cichy, R. M., & Woolrich, M. W. (2021) · Cerebral Cortex · doi:10.1093/cercor/bhab189 · Cited by 15 · see citing works
“modulate neural responses (Podvalny et al. 2019; McCormick et al. 2020), it is clear that part of the large variability observed in the neural responses could be explained by the ongoing, largescale network activity and concurrent phase trajectories, and that accounting by these factors could arguably improve the capa…”
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Midbrain encodes sound detection behavior without auditory cortexPreprint
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Recurrent network interactions explain tectal response variability and experience-dependent behavior
Zylbertal, A., & Bianco, I. H. (2023) · eLife · doi:10.7554/elife.78381 · Cited by 11 · see citing works
Contribution of membrane-associated oscillators to biological timing at different timescales
Stengl, M., & Schneider, A. C. (2024) · Frontiers in Physiology · doi:10.3389/fphys.2023.1243455 · Cited by 15 · see citing works
“As foundation of robust, autonomous neuronal functions multiple mechanisms of homeostatic plasticity at the level of single neurons as well as at the level of neuronal ensembles maintain a speci fi c dynamic setpoint measurable as stable spike frequency (McCormick et al., 2020).”
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“, 2014; Mostame and Sadaghiani, 2020) and neuromodulation (McCormick et al., 2020; Pfeffer et al., 2021; van den Brink et al., 2018; van den Brink et al., 2019).”
BackgroundInput-Independent Homeostasis of Developing Thalamocortical Activity
Riyahi, P., Phillips, M. A., & Colonnese, M. T. (2021) · eNeuro · doi:10.1523/eneuro.0184-21.2021 · Cited by 11 · see citing works
Ngomba, R. T., Lüttjohann, A., Dexter, A., Ray, S., & van Luijtelaar, G. (2023) · Cells · doi:10.3390/cells12131761 · Cited by 10 · see citing works
“Different brain regions, including the neocortex, hippocampus, thalamus, and brainstem, are involved in the neuronal activities during the various sleep states [14].”
BackgroundBrain network dynamics predict moments of surprise across contexts
Zhang, Z., & Rosenberg, M. D. (2024) · Nature Human Behaviour · doi:10.1038/s41562-024-02017-0 · Cited by 10 · see citing works
“This approach offers new perspectives for analysing data from existing cognitive paradigms and possibilities for identifying signatures of cognitive states fluctuations from large-scale brain networks 69–71 .”
Excite the unexcitable: engineering cells and redox signaling for targeted bioelectronic control
Wang, S., Aljirafi, F. O., Payne, G. F., & Bentley, W. E. (2023) · Current Opinion in Biotechnology · doi:10.1016/j.copbio.2023.103052 · Cited by 10 · see citing works
- Authors
- McCormick, D. A., Nestvogel, D. B., & He, B. J.
- Affiliations
- University of Oregon (US)
- NYU Langone Health (US)
- Corresponding author
- McCormick, David A.
- Venue
- Annual Review of Neuroscience
- Publisher
- Annual Reviews
- Published
- 2020-07-08
- Volume
- 43
- Issue
- 1
- Pages
- 391-415
- ISSN
- 0147-006X (ISSN-L)
- 1545-4126
- PMC
- PMC12237593
- OpenAlex
- W3014163957
- Cited by
- 303
- References
- 167
- Citations by year
- 2026: 40 · 2025: 60 · 2024: 72 · 2023: 48 · 2022: 41 · 2021: 36 · 2020: 6
- FWCI
- 4.43
- Type
- journal article
- Language
- en
- Open access
- green
- www.ncbi.nlm.nih.gov
- Locations
- Annual Review of Neuroscience (publishedVersion)
- PubMed (publishedVersion)
- PubMed Central (acceptedVersion)
- MPG.PuRe (Max Planck Society) (submittedVersion)
- Funders
- National Institute on Deafness and Other Communication Disorders (R01DC015803)
- National Institute of Neurological Disorders and Stroke (R35NS097287)
- Topics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Sleep and Wakefulness Research
- Life Sciences › Neuroscience › Cognitive Neuroscience › Neural dynamics and brain function
- Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
- Keywords
- brain states, neural state transitions, behavioral state transitions, neuromodulation, cortico-cortical pathways, neural variability, cognitive states
- MeSH terms
- Animals, Behavior, Brain, Cerebral Cortex, Humans, Nerve Net, Neural Pathways, Neurons
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
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