LSDPsilocybinMescaline

Recent advances in the neuropsychopharmacology of serotonergic hallucinogens

This study (2015) reviews the evidence on the neuropsychopharmacology of such substances as LSD, psilocybin, and mescaline.

Authors

  • Halberstadt, A. L.

Published

Behavioural Brain Research
meta Study

Abstract

Serotonergic hallucinogens, such as (+)-lysergic acid diethylamide, psilocybin, and mescaline, are somewhat enigmatic substances. Although these drugs are derived from multiple chemical families, they all produce remarkably similar effects in animals and humans, and they show cross-tolerance. This article reviews the evidence demonstrating the serotonin 5-HT2A receptor is the primary site of hallucinogen action. The 5-HT2A receptor is responsible for mediating the effects of hallucinogens in human subjects, as well as in animal behavioral paradigms such as drug discrimination, head twitch response, prepulse inhibition of startle, exploratory behavior, and interval timing. Many recent clinical trials have yielded important new findings regarding the psychopharmacology of these substances. Furthermore, the use of modern imaging and electrophysiological techniques is beginning to help unravel how hallucinogens work in the brain. Evidence is also emerging that hallucinogens may possess therapeutic efficacy.

Unlocked with Blossom Pro

Research Summary of 'Recent advances in the neuropsychopharmacology of serotonergic hallucinogens'

Introduction

Serotonergic hallucinogens — encompassing two main structural classes, the phenylalkylamines and the indoleamines — produce profound alterations in perception, thought, and mood and have been used by humans for thousands of years, though scientific investigation began only in the late nineteenth century. Defining hallucinogens as a pharmacological class proved contentious: standard definitions based on altered consciousness without delirium or addiction were insufficiently specific, leading to the proposal that, in addition to these properties, hallucinogens should bind to the 5-HT2A receptor and produce full substitution for the prototypical agent DOM in drug discrimination paradigms. Despite their structural heterogeneity, phenylalkylamine and indoleamine hallucinogens produce virtually indistinguishable subjective effects, show cross-tolerance with one another, and share a unifying pharmacological mechanism through 5-HT2A receptor activation. This review aimed to synthesise advances in understanding the neuropsychopharmacology of serotonergic hallucinogens, covering receptor pharmacology, validated animal behavioural models, tolerance and cross-tolerance, evidence from human experimental studies, and the role of the prefrontal cortex and its interactions with subcortical structures as a primary site of hallucinogenic drug action.

Methods

This is a comprehensive narrative review that synthesised the preclinical and human experimental literature on serotonergic hallucinogens without formal inclusion/exclusion criteria or quantitative meta-analytic methods. The author covered receptor binding and functional pharmacology — including 5-HT2A and 5-HT2C subtype selectivity — drug discrimination studies using operant conditioning paradigms in rodents, the head twitch response (HTR) assay, prepulse inhibition of startle (PPI), interval timing paradigms, and locomotor activity measures. Human experimental evidence was drawn from challenge studies examining cross-tolerance and subjective effects across compounds. Neurobiological coverage focused on 5-HT2A receptor expression patterns across cortical layers and brain regions, electrophysiological recordings from prefrontal cortex pyramidal neurons in vitro and in vivo, and the organisation of cortico-striato-thalamo-cortical (CSTC) loop circuitry as a substrate for hallucinogen action.

Results

Cross-tolerance studies confirmed that phenylalkylamine and indoleamine hallucinogens mutually substitute in drug discrimination paradigms and produce equivalent blunting of subjective effects following repeated dosing — evidence consistent with a shared receptor mechanism. Multiple converging lines of evidence from in vitro and in vivo work identified 5-HT2A receptor activation as the unitary mechanism responsible for hallucinogenesis: selective 5-HT2A antagonists block hallucinogenic behavioural effects in animals and subjective effects in humans, and highly selective phenylisopropylamine hallucinogens such as DOM produce full substitution for indoleamines in drug discrimination. Electrophysiological investigation identified the prefrontal cortex as a primary site of hallucinogen action. The 5-HT2A receptor is expressed densely in lamina V of the PFC on pyramidal neurons and parvalbumin-positive interneurons. Systemic hallucinogen administration in rats produced a net excitatory effect on PFC pyramidal neurons, with individual cells either excited (38–53%), inhibited (27–35%), or unresponsive. These PFC effects are thought to propagate through CSTC loops involving thalamo-cortical and striato-thalamic circuits to generate the broader pattern of perceptual and cognitive alterations characteristic of hallucinogen action.

Discussion

Convergence of pharmacological, electrophysiological, and human experimental evidence leaves little scientific doubt that 5-HT2A receptor activation is the primary mechanism of classical hallucinogenesis, with the prefrontal cortex — and its dense laminar expression of the receptor on pyramidal neurons — constituting a critical neuroanatomical site of drug action. The net excitatory effect of hallucinogens on PFC pyramidal cells, propagated through CSTC circuitry, offers a plausible neural basis for the disorganised and amplified pattern of cortical activity associated with altered consciousness and perceptual distortion. The resumption of human research with hallucinogens — including brain imaging studies using fMRI, PET, and MEG — has provided additional support for animal and in vitro evidence, and is beginning to elucidate how receptor-level pharmacology translates into circuit-level and behavioural effects. The author identifies understanding of functional selectivity at the 5-HT2A receptor — specifically the relative contributions of G-protein and β-arrestin signalling pathways — as an important avenue for dissociating perceptual from therapeutic effects and for the rational design of functionally selective compounds.

Conclusion

The neuropsychopharmacology of serotonergic hallucinogens is increasingly well characterised: both phenylalkylamine and indoleamine classes produce their subjective and behavioural effects primarily through 5-HT2A receptor activation, with the prefrontal cortex and cortico-striato-thalamo-cortical circuits serving as the principal neuroanatomical substrates. Validated animal models — including drug discrimination, head twitch response, and prepulse inhibition paradigms — have provided essential tools for dissecting receptor mechanisms, and the resumption of controlled human studies is beginning to translate these preclinical insights into neuroimaging and clinical findings. Continued investigation of functional selectivity at the 5-HT2A receptor and the circuit-level consequences of PFC activation holds promise for developing a more complete account of hallucinogen pharmacology and informing the design of novel therapeutic agents.

Study Details

References (33)

Papers cited by this study that are also in Blossom

Multiple receptors contribute to the behavioral effects of indoleamine hallucinogens

Geyer, M. A., Halberstadt, A. L. · Neuropharmacology (2011)

Dose-related effects of salvinorin A in humans: dissociative, hallucinogenic, and memory effects

Griffiths, R. R., Johnson, M. W., MacLean, K. A. et al. · Psychopharmacology (2012)

Psilocybin can occasion mystical-type experiences having substantial and sustained personal meaning and spiritual significance

Griffiths, R. R., Jesse, R., McCann, U. D. et al. · Journal of Psychopharmacology (2006)

187 cited
Psilocybin occasioned mystical-type experiences: immediate and persisting dose-related effects

Griffiths, R. R., Jesse, R., Johnson, M. W. et al. · Psychopharmacology (2011)

Psychometric evaluation of the altered states of consciousness rating scale (OAV)

Gamma, A., Studerus, E., Vollenweider, F. X. · PLOS ONE (2010)

Show all 33 references
Psilocybin induces schizophrenia-like psychosis in humans via a serotonin-2 agonist action

Bäbler, A., Hell, D., Vogel, H. et al. · NeuroReport (1998)

Using Psilocybin to Investigate the Relationship between Attention, Working Memory, and the Serotonin 1A and 2A Receptors

Burr, D. C., Carter, O., Pettigrew, J. D. et al. · Journal of Cognitive Neuroscience (2006)

Psilocybin links binocular rivalry switch rate to attention and subjective arousal levels in humans

Carter, O., Hasler, F., Liu, G. B. et al. · Psychopharmacology (2007)

Psilocybin-Induced Deficits in Automatic and Controlled Inhibition are Attenuated by Ketanserin in Healthy Human Volunteers

Geyer, M. A., Kometer, M., Quednow, B. B. et al. · Neuropsychopharmacology (2011)

Functional selectivity of hallucinogenic phenethylamine and phenylisopropylamine derivatives at human 5-hydroxytryptamine (5-HT) 2A and 5-HT2C receptors

Berg, K. A., Cassels, B. K., Clarke, W. P. et al. · Journal of Pharmacology and Experimental Therapeutics (2007)

Behavioral, neurochemical and pharmaco-EEG profiles of the psychedelic drug 4-bromo-2, 5-dimethoxyphenethylamine (2C-B) in rats

Balíková, M., Brunovský, M., Bubeníková-Valešová, V. et al. · Psychopharmacology (2012)

Effects of mescaline and lysergic acid (d-LSD-25)

Cattell, J. P., Hoch, P. H., Pennes, H. H. · American Journal of Psychiatry (1987)

Influence of Lysergic Acid Diethylamide (LSD-25) on Subjective Time

Aronson, H., Klee, G. D., Silverstein, A. B. · JAMA Psychiatry (1980)

Effects of psilocybin on time perception and temporal control of behaviour in humans

Cahn, B. R., Carter, O., Flohr, H. et al. · Journal of Psychopharmacology (2007)

212 cited
The psychedelic model of schizophrenia: the case of N,N-dimethyltryptamine

Gillin, J. C., Kaplan, J., Stillman, R. et al. · American Journal of Psychiatry (1976)

Broadband Cortical Desynchronization Underlies the Human Psychedelic State

Bolstridge, M., Brookes, M. J., Carhart-Harris, R. L. et al. · Journal of Neuroscience (2013)

427 cited
Topographic pharmaco-EEG mapping of the effects of the South American psychoactive beverage ayahuasca in healthy volunteers

Anderer, P., Barbanoj, M. J., Jané, F. et al. · British Journal of Clinical Pharmacology (2002)

Neural correlates of the psychedelic state as determined by fMRI studies with psilocybin

Carhart-Harris, R. L., Colasanti, A., Erritzoe, D. et al. · PNAS (2012)

Implications for psychedelic-assisted psychotherapy: functional magnetic resonance imaging study with psilocybin

Abbasi, N., Bargiotas, T., Carhart-Harris, R. L. et al. · British Journal of Psychiatry (2018)

202 cited
Psilocybin-induced decrease in amygdala reactivity correlates with enhanced positive mood in healthy volunteers

Bosch, O. G., Kraehenmann, R., Pokorny, T. et al. · Biological Psychiatry (2015)

The 5-HT2A/1A Agonist Psilocybin Disrupts Modal Object Completion Associated with Visual Hallucinations

Andel, D., Cahn, B. R., Carter, O. et al. · Biological Psychiatry (2011)

74 cited
A review of the clinical effects of psychotomimetic agents

Osmond, H. · Annals of the New York Academy of Sciences (2010)

296 cited
Safety and efficacy of lysergic acid diethylamide-assisted psychotherapy for anxiety associated with life-threatening diseases

Brenneisen, R., Doblin, R., Gasser, P. et al. · Journal of Nervous and Mental Disease (2014)

594 cited

Cited By (72)

Papers in Blossom that reference this study

Psilocybin alters visual contextual computations

Aqil, M., de Hollander, G., Dumoulin, S. O. et al. · Biorxiv (2025)

Membrane Permeation of Psychedelic Tryptamines by Dynamic Simulations

Agnorelli, C., Erritzoe, D., Fagiolini, A. et al. · Biochemistry (2024)

Synthetic surprise as the foundation of the psychedelic experience

De Filippo, R., Schmitz, D. · Neuroscience and Biobehavioral Reviews (2024)

Identification of 5-HT 2A Receptor Signaling Pathways Responsible for Psychedelic Potential

Anderson, E. I., Bock, H. A., Bonniwell, E. M. et al. · Biorxiv (2023)

Impact of a Naturalistic Psychedelic Experience on Smoking: A Retrospective Survey

Benyamina, A., Fauvel, B., Piolino, P. et al. · Journal of Psychoactive Drugs (2023)

Dose-response relationships of LSD-induced subjective experiences in humans

Hirschfeld, T., Majic, T., Prugger, J. et al. · Neuropsychopharmacology (2023)

A non-hallucinogenic LSD analog with therapeutic potential for mood disorders

Abizaid, A., Aguilar-Valles, A., Arsenault, E. et al. · Cell Reports (2023)

102 cited
Show all 72 papers
Neural Mechanisms and Psychology of Psychedelic Ego Dissolution

Egan, G. F., Friston, K. J., Razi, A. et al. · Pharmacological Reviews (2022)

Classic psychedelics and alcohol use disorders: A systematic review of human and animal studies

Bühler, K. M., Calleja-Conde, J., Echeverry-Alzate, V. et al. · Addiction Biology (2022)

Psychedelic Microdosing, Mindfulness, and Anxiety: A Cross-Sectional Mediation Study

Hartong, V., van Emmerik, A. · Journal of Psychoactive Drugs (2022)

Metabolomics and integrated network analysis reveal roles of endocannabinoids and large neutral amino acid balance in the ayahuasca experience

Busquets-Garcia, A., de la Torre, R., Gomez-Gomez, A. et al. · Biomedicine & Pharmacotherapy (2022)

Classic Psychedelic Drugs: Update on Biological Mechanisms

Smallridge, J. W., Vollenweider, F. X. · Pharmacopsychiatry (2022)

5-HT2CR Is as Important as 5-HT2AR in Inducing Hallucinogenic Effects in Serotonergic Compounds

Buctot, D., Cheong, J. H., Custodio, R. J. et al. · SSRN (2022)

Toward a positive psychology of psychoactive drug use

Arnaud, K. O. S. · Drugs Education Prevention and Policy (2021)

Psilocybin as a Novel Pharmacotherapy for Treatment-Refractory Anorexia Nervosa

Aouad, P., Maguire, S., McGregor, I. S. et al. · OBM Neurobiology (2021)

Hallucinogenic/psychedelic 5HT2A receptor agonists as rapid antidepressant therapeutics: Evidence and mechanisms of action

Baker, G., Dos Santos, R. G., Dursun, S. et al. · Journal of Psychopharmacology (2021)

Psilocin acutely disrupts sleep and affects local but not global sleep homeostasis in laboratory mice

Bannerman, D., Blanco-Duque, C., Breant, B. et al. · Translational Psychiatry (2021)

Investigation of the Structure-Activity Relationships of Psilocybin Analogues

Anderson, E. I., Brandt, S. D., Chapman, S. J. et al. · ACS Pharmacology and Translational Science (2020)

Psychedelic drugs: neurobiology and potential for treatment of psychiatric disorders

Preller, K. H., Vollenweider, F. X. · Nature Reviews Neuroscience (2020)

73 cited
Microdosing psychedelics: Demographics, practices, and psychiatric comorbidities.

Anderson, T., Dinh-Williams, L., Hapke, E. et al. · Journal of Psychopharmacology (2020)

55 cited
Psychedelics and Psychedelic-Assisted Psychotherapy

Carpenter, L. L., Kalin, N. H., McDonald, W. et al. · American Journal of Psychiatry (2020)

Spectral signatures of serotonergic psychedelics and glutamatergic dissociatives

Carhart-Harris, R. L., Muthukumaraswamy, S., Nutt, D. J. et al. · NeuroImage (2019)

51 cited
Metabolism of lysergic acid diethylamide (LSD): an update

Libânio Osório Marta, R. F. · Drug Metabolism Reviews (2019)

The effects of microdose LSD on time perception: a randomised, double-blind, placebo-controlled trial

Family, N., Luke, D. P., Polychroni, N. et al. · Psychopharmacology (2018)

Whole-brain multimodal neuroimaging model using serotonin receptor maps explains non-linear functional effects of LSD

Cabral, J., Carhart-Harris, R. L., Cruzat, J. et al. · Current Biology (2018)

Psychedelics, meditation, and self-consciousness

Berkovich-Ohana, A., Carhart-Harris, R. L., Milliere, R. et al. · Frontiers in Psychology (2018)

Dark Classics in Chemical Neuroscience: N, N-Dimethyltryptamine (DMT)

Cameron, L. P. · ACS Chemical Neuroscience (2018)

Unifying theories of psychedelic drug effects

Swanson, L. R. · Frontiers in Pharmacology (2018)

Serotonergic hallucinogens in the treatment of anxiety and depression in patients suffering from a life-threatening disease: A systematic review

Gasser, P., Gutwinski, S., Hermle, L. et al. · Progress in Neuro-Psychopharmacology and Biological Psychiatry (2018)

Hallucinogens and Serotonin 5-HT2A Receptor-Mediated Signaling Pathways

González-Maeso, J., López-Giménez, J. F. · Behavioral Neurobiology of Psychedelic Drugs (2018)

Serotonin and brain function: a tale of two receptors

Carhart-Harris, R. L., Nutt, D. J. · Journal of Psychopharmacology (2017)

Clinical Applications of Hallucinogens: A Review

Addy, P. H., Garcia-Romeu, A., Kersgaard, B. · Experimental and Clinical Psychopharmacology (2017)

Modern clinical research on LSD

Liechti, M. E. · Neuropsychopharmacology (2017)

Increased spontaneous MEG signal diversity for psychoactive doses of ketamine, LSD and psilocybin

Barrett, A. B., Carhart-Harris, R. L., Muthukumaraswamy, S. et al. · Scientific Reports (2017)

Dreamlike effects of LSD on waking imagery in humans depend on serotonin 2A receptor activation

Kraehenmann, R., Pokorny, D., Pokorny, T. et al. · Psychopharmacology (2017)

138 cited
5-HT2A Agonists: A Novel Therapy for Functional Neurological Disorders?

Bryson, A., Carter, O., Kanaan, R. A. et al. · International Journal of Neuropsychopharmacology (2017)

18 cited
The fabric of meaning and subjective effects in LSD-induced states depend on serotonin 2A receptor activation

Herdener, M., Kraehenmann, R., Liechti, M. E. et al. · Current Biology (2017)

Pharmacology and Toxicology of N-Benzylphenethylamine (“NBOMe”) Hallucinogens

Halberstadt, A. L. · Current Topics in Behavioral Neurosciences (2017)

Phenomenology, Structure, and Dynamic of Psychedelic States

Preller, K. H., Vollenweider, F. X. · Behavioral Neurobiology of Psychedelic Drugs (2016)

The Challenging Experience Questionnaire: Characterization of challenging experiences with psilocybin mushrooms

Barrett, F. S., Bradstreet, M. P., Griffiths, R. R. et al. · Journal of Psychopharmacology (2016)

The serotonin 5-HT2C receptor and the non-addictive nature of classic hallucinogens

Cabral, J., Canal, C. E., Murnane, K. S. · Journal of Psychopharmacology (2016)

Classical hallucinogens and neuroimaging: A systematic review of human studies: hallucinogens and neuroimaging

Crippa, J. A., Dos Santos, R. G., Hallak, J. E. et al. · Neuroscience and Biobehavioral Reviews (2016)

Development of a Psychotherapeutic Model for Psilocybin-Assisted Treatment of Alcoholism

Bogenschutz, M. P., Forcehimes, A. A. · Journal of Humanistic Psychology (2016)

Receptor interaction profiles of novel psychoactive tryptamines compared with classic hallucinogens

Hoener, M. C., Liechti, M. E., Moning, O. D. et al. · European Neuropsychopharmacology (2016)

Treating Addiction: Perspectives from EEG and Imaging Studies on Psychedelics

de Araujo, D. B., Tófoli, L.F. · International Review of Neurobiology (2016)

LSD alters eyes-closed functional connectivity within the early visual cortex in a retinotopic fashion

Carhart-Harris, R. L., Feilding, A., Kaelen, M. et al. · Human Brain Mapping (2016)

Neural correlates of the LSD experience revealed by multimodal neuroimaging

Bolstridge, M., Carhart-Harris, R. L., Curran, H. V. et al. · PNAS (2016)

Psychedelics

Nichols, D. E. · Pharmacological Reviews (2016)

New World Tryptamine Hallucinogens and the Neuroscience of Ayahuasca

McKenna, D., Riba, J. · Current Topics in Behavioral Neurosciences (2016)

Serotonergic Hallucinogen-Induced Visual Perceptual Alterations

Kometer, M., Vollenweider, F. X. · Behavioral Neurobiology of Psychedelic Drugs (2016)

Psychedelics not linked to mental health problems or suicidal behavior: A population study.

Johansen, P. Ø., Krebs, T. S. · Journal of Psychopharmacology (2015)

Your Library