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Vol. 53, Issue 1, 107-118, March 2001

International Union of Pharmacology. XXIV. Current Status of the Nomenclature and Properties of P2X Receptors and Their Subunits

Baljit S. Khakh1, Geoffrey Burnstock, Charles Kennedy, Brian F. King, R. Alan North, Philippe Séguéla, Mark Voigt and Patrick P. A. Humphrey1

Division of Biology 156-29, California Institute of Technology, Pasadena, California (B.S.K.); Autonomic Neuroscience Institute, Royal Free Hospital School of Medicine, London, United Kingdom (G.B., B.F.K.); Department of Physiology and Pharmacology, University of Strathclyde, Royal College, Glasgow, United Kingdom (C.K.); Institute of Molecular Physiology, University of Sheffield, Western Bank, Sheffield, United Kingdom (R.A.N.); Department of Neurology and Neurosurgery, Montreal Neurological Institute, McGill University, Montreal, Quebec, Canada (P.S.); Pharmacological and Physiological Sciences, St. Louis University School of Medicine, St. Louis, Missouri (M.V.); and Glaxo Institute of Applied Pharmacology, University of Cambridge, Department of Pharmacology, Cambridge, United Kingdom (P.P.A.H.)

I. Introduction
II. Overall P2X Subunit Topology
III. Electrophysiological Properties of P2X Receptors
IV. Functional Properties of Homomeric Receptors
    A. P2X1
    B. P2X2
    C. P2X3
    D. P2X4
    E. P2X5
    F. P2X6
    G. P2X7
V. Functional Properties of Heteromeric Receptors
VI. Native Receptors in Whole Tissues
    A. Studies in Vitro
    B. Studies in Vivo
VII. Summary
Acknowledgments
References

ATP acts as a humoral mediator to control cell function extracellularly. The receptors that mediate the actions of ATP belong to two classes, the metabotropic P2Y receptors and the transmitter-gated, ion channel P2X receptors. This review describes the structure, distribution, function, and ligand recognition characteristics of P2X receptors, which comprise seven distinct subunits that can function as both homo- and hetero- polymers. The pharmacology of P2X receptors is complicated by marked differences between species orthologues. The current nomenclature is based largely on recombinant receptor studies and detailed knowledge of endogenous P2X receptors in native tissues is limited because of lack of good selective agonists and antagonists for each receptor type.

1 Address for correspondence: Baljit S. Khakh, Division of Neurobiology, MRC Laboratory of Molecular Biology, Hills Rd., Cambridge, CB2 2QH, UK. E-mail: bsk{at}mrc-lmb.cam.ac.uk or Patrick P. A. Humphrey, Glaxo Institute of Applied Pharmacology, University of Cambridge, Department of Pharmacology, Tennis Court Rd., Cambridge, CB2 1QJ UK. E-mail: ppah0562{at}glaxowellcome.co.uk

0031-6997/01/5301-0107$03.00/0
PHARMACOLOGICAL REVIEWS
Copyright © 2001 by The American Society for Pharmacology and Experimental Therapeutics


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R. Schafer, F. Sedehizade, T. Welte, and G. Reiser
ATP- and UTP-activated P2Y receptors differently regulate proliferation of human lung epithelial tumor cells
Am J Physiol Lung Cell Mol Physiol, August 1, 2003; 285(2): L376 - L385.
[Abstract] [Full Text] [PDF]

Home page
 Ann. Thorac. Surg.Home page
C. Borna, L. Wang, T. Gudbjartsson, L. Karlsson, S. Jern, M. Malmsjo, and D. Erlinge
Contractions in human coronary bypass vessels stimulated by extracellular nucleotides
Ann. Thorac. Surg., July 1, 2003; 76(1): 50 - 57.
[Abstract] [Full Text] [PDF]

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 JGPHome page
A. Skorinkin, A. Nistri, and R. Giniatullin
Bimodal Action of Protons on ATP Currents of Rat PC12 Cells
J. Gen. Physiol., June 30, 2003; 122(1): 33 - 44.
[Abstract] [Full Text] [PDF]

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 J. Neurophysiol.Home page
K. Tsuzuki, A. Ase, P. Seguela, T. Nakatsuka, C.-Y. Wang, J.-X. She, and J. G. Gu
TNP-ATP-Resistant P2X Ionic Current on the Central Terminals and Somata of Rat Primary Sensory Neurons
J Neurophysiol, June 1, 2003; 89(6): 3235 - 3242.
[Abstract] [Full Text] [PDF]

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 J. Neurophysiol.Home page
T. Nakatsuka, K. Tsuzuki, J. X. Ling, H. Sonobe, and J. G. Gu
Distinct Roles of P2X Receptors in Modulating Glutamate Release at Different Primary Sensory Synapses in Rat Spinal Cord
J Neurophysiol, June 1, 2003; 89(6): 3243 - 3252.
[Abstract] [Full Text] [PDF]

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 J. Leukoc. Biol.Home page
P. Bradding, Y. Okayama, N. Kambe, and H. Saito
Ion channel gene expression in human lung, skin, and cord blood-derived mast cells
J. Leukoc. Biol., May 1, 2003; 73(5): 614 - 620.
[Abstract] [Full Text] [PDF]

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 J. Biol. Chem.Home page
M.-L. He, H. Zemkova, and S. S. Stojilkovic
Dependence of Purinergic P2X Receptor Activity on Ectodomain Structure
J. Biol. Chem., March 14, 2003; 278(12): 10182 - 10188.
[Abstract] [Full Text] [PDF]

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 PhysiologyHome page
I. Novak
ATP as a Signaling Molecule: the Exocrine Focus
Physiology, February 1, 2003; 18(1): 12 - 17.
[Abstract] [Full Text] [PDF]

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 J. Exp. Biol.Home page
C. R. Wood and T. M. Hennessey
PPNDS is an agonist, not an antagonist, for the ATP receptor of Paramecium
J. Exp. Biol., February 1, 2003; 206(3): 627 - 636.
[Abstract] [Full Text] [PDF]

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 Am. J. Physiol. Gastrointest. Liver Physiol.Home page
C. Barajas-Lopez, L. M. Montano, and R. Espinosa-Luna
Inhibitory interactions between 5-HT3 and P2X channels in submucosal neurons
Am J Physiol Gastrointest Liver Physiol, December 1, 2002; 283(6): G1238 - G1248.
[Abstract] [Full Text] [PDF]

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 J. Biol. Chem.Home page
T.-a. Koshimizu, S. Ueno, A. Tanoue, N. Yanagihara, S. S. Stojilkovic, and G. Tsujimoto
Heteromultimerization Modulates P2X Receptor Functions through Participating Extracellular and C-terminal Subdomains
J. Biol. Chem., November 27, 2002; 277(49): 46891 - 46899.
[Abstract] [Full Text] [PDF]

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 J. Biol. Chem.Home page
E. V. Gerasimovskaya, S. Ahmad, C. W. White, P. L. Jones, T. C. Carpenter, and K. R. Stenmark
Extracellular ATP Is an Autocrine/Paracrine Regulator of Hypoxia-induced Adventitial Fibroblast Growth. SIGNALING THROUGH EXTRACELLULAR SIGNAL-REGULATED KINASE-1/2 AND THE Egr-1 TRANSCRIPTION FACTOR
J. Biol. Chem., November 15, 2002; 277(47): 44638 - 44650.
[Abstract] [Full Text] [PDF]

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 J. Neurophysiol.Home page
B. Hu, C. Y. Chiang, J. W. Hu, J. O. Dostrovsky, and B. J. Sessle
P2X Receptors in Trigeminal Subnucleus Caudalis Modulate Central Sensitization in Trigeminal Subnucleus Oralis
J Neurophysiol, October 1, 2002; 88(4): 1614 - 1624.
[Abstract] [Full Text] [PDF]

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 Mol. Pharmacol.Home page
S. S. Wildman, S. G. Brown, M. Rahman, C. A. Noel, L. Churchill, G. Burnstock, R. J. Unwin, and B. F. King
Sensitization by Extracellular Ca2+ of Rat P2X5 Receptor and Its Pharmacological Properties Compared with Rat P2X1.
Mol. Pharmacol., October 1, 2002; 62(4): 957 - 966.
[Abstract] [Full Text] [PDF]

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 J. Biol. Chem.Home page
S. J. Royle, L. K. Bobanovic', and R. D. Murrell-Lagnado
Identification of a Non-canonical Tyrosine-based Endocytic Motif in an Ionotropic Receptor
J. Biol. Chem., September 13, 2002; 277(38): 35378 - 35385.
[Abstract] [Full Text] [PDF]

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 J. Neurosci.Home page
P. P. Bertrand and J. C. Bornstein
ATP as a Putative Sensory Mediator: Activation of Intrinsic Sensory Neurons of the Myenteric Plexus via P2X Receptors
J. Neurosci., June 15, 2002; 22(12): 4767 - 4775.
[Abstract] [Full Text] [PDF]

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 J. Neurosci.Home page
L. K. Bobanovic, S. J. Royle, and R. D. Murrell-Lagnado
P2X Receptor Trafficking in Neurons Is Subunit Specific
J. Neurosci., June 15, 2002; 22(12): 4814 - 4824.
[Abstract] [Full Text] [PDF]

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 J. Biol. Chem.Home page
B. Hu, C. Senkler, A. Yang, F. Soto, and B. T. Liang
P2X4 Receptor Is a Glycosylated Cardiac Receptor Mediating a Positive Inotropic Response to ATP
J. Biol. Chem., May 3, 2002; 277(18): 15752 - 15757.
[Abstract] [Full Text] [PDF]

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 J. Neurosci.Home page
T. Nakatsuka, H. Furue, M. Yoshimura, and J. G. Gu
Activation of Central Terminal Vanilloid Receptor-1 Receptors and alpha beta -Methylene-ATP-Sensitive P2X Receptors Reveals a Converged Synaptic Activity onto the Deep Dorsal Horn Neurons of the Spinal Cord
J. Neurosci., February 15, 2002; 22(4): 1228 - 1237.
[Abstract] [Full Text] [PDF]

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 BloodHome page
E. Adinolfi, L. Melchiorri, S. Falzoni, P. Chiozzi, A. Morelli, A. Tieghi, A. Cuneo, G. Castoldi, F. Di Virgilio, and O. R. Baricordi
P2X7 receptor expression in evolutive and indolent forms of chronic B lymphocytic leukemia
Blood, January 15, 2002; 99(2): 706 - 708.
[Abstract] [Full Text] [PDF]

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 J. Neurosci.Home page
J. H. Lee, T. Chiba, and D. C. Marcus
P2X2 Receptor Mediates Stimulation of Parasensory Cation Absorption by Cochlear Outer Sulcus Cells and Vestibular Transitional Cells
J. Neurosci., December 1, 2001; 21(23): 9168 - 9174.
[Abstract] [Full Text] [PDF]

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 Pharmacol. Rev.Home page
B. A. Chizh and P. Illes
P2X Receptors and Nociception
Pharmacol. Rev., December 1, 2001; 53(4): 553 - 568.
[Abstract] [Full Text] [PDF]

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 J. Physiol.Home page
I.-S. Jang, J. S. Rhee, H. Kubota, N. Akaike, and N. Akaike
Developmental changes in P2X purinoceptors on glycinergic presynaptic nerve terminals projecting to rat substantia gelatinosa neurones
J. Physiol., October 15, 2001; 536(2): 505 - 519.
[Abstract] [Full Text] [PDF]

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 J. Biol. Chem.Home page
L. N. Naemsch, S. J. Dixon, and S. M. Sims
Activity-dependent Development of P2X7 Current and Ca2+ Entry in Rabbit Osteoclasts
J. Biol. Chem., October 12, 2001; 276(42): 39107 - 39114.
[Abstract] [Full Text] [PDF]

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 J. Neurosci.Home page
T. Nakatsuka and J. G. Gu
ATP P2X Receptor-Mediated Enhancement of Glutamate Release and Evoked EPSCs in Dorsal Horn Neurons of the Rat Spinal Cord
J. Neurosci., September 1, 2001; 21(17): 6522 - 6531.
[Abstract] [Full Text] [PDF]

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 J. Neurosci.Home page
W. R. Haines, M. M. Voigt, K. Migita, G. E. Torres, and T. M. Egan
On the Contribution of the First Transmembrane Domain to Whole-Cell Current through an ATP-Gated Ionotropic P2X Receptor
J. Neurosci., August 15, 2001; 21(16): 5885 - 5892.
[Abstract] [Full Text] [PDF]

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 Proc. Natl. Acad. Sci. USAHome page
B. S. Khakh, W. B. Smith, C.-S. Chiu, D. Ju, N. Davidson, and H. A. Lester
Activation-dependent changes in receptor distribution and dendritic morphology in hippocampal neurons expressing P2X2-green fluorescent protein receptors
PNAS, April 5, 2001; (2001) 81089198.
[Abstract] [Full Text]

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 J. Biol. Chem.Home page
K. Migita, W. R. Haines, M. M. Voigt, and T. M. Egan
Polar Residues of the Second Transmembrane Domain Influence Cation Permeability of the ATP-gated P2X2 Receptor
J. Biol. Chem., August 10, 2001; 276(33): 30934 - 30941.
[Abstract] [Full Text] [PDF]

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 J. Biol. Chem.Home page
W. R. Haines, K. Migita, J. A. Cox, T. M. Egan, and M. M. Voigt
The First Transmembrane Domain of the P2X Receptor Subunit Participates in the Agonist-induced Gating of the Channel
J. Biol. Chem., August 24, 2001; 276(35): 32793 - 32798.
[Abstract] [Full Text] [PDF]

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 Proc. Natl. Acad. Sci. USAHome page
B. S. Khakh, W. B. Smith, C.-S. Chiu, D. Ju, N. Davidson, and H. A. Lester
Activation-dependent changes in receptor distribution and dendritic morphology in hippocampal neurons expressing P2X2-green fluorescent protein receptors
PNAS, April 24, 2001; 98(9): 5288 - 5293.
[Abstract] [Full Text] [PDF]


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