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Review ArticleReview

International Union of Pharmacology. XXVII. Classification of Cannabinoid Receptors

A. C. Howlett, F. Barth, T. I. Bonner, G. Cabral, P. Casellas, W. A. Devane, C. C. Felder, M. Herkenham, K. Mackie, B. R. Martin, R. Mechoulam and R. G. Pertwee
Pharmacological Reviews June 2002, 54 (2) 161-202; DOI: https://doi.org/10.1124/pr.54.2.161
A. C. Howlett
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F. Barth
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T. I. Bonner
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G. Cabral
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P. Casellas
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W. A. Devane
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C. C. Felder
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M. Herkenham
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K. Mackie
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B. R. Martin
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R. Mechoulam
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R. G. Pertwee
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  • Figure 1
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    Figure 1

    The structures of four constituents of cannabis: Δ9-THC, Δ8-THC, cannabinol, and cannabidiol.

  • Figure 2
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    Figure 2

    The structures of the synthetic classical cannabinoid receptor agonists, HU-210 and desacetyl-l-nantradol, and of HU-211, the (+)-enantiomer of HU-210.

  • Figure 3
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    Figure 3

    The structures of the CB2-selective cannabinoid receptor agonists, HU-308, L-759633, L-759656, JWH-133, JWH-139, and JWH-051.

  • Figure 4
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    Figure 4

    The structures of the (−)-enantiomers of three nonclassical cannabinoid receptor agonists: CP55940, CP47497, and CP55244.

  • Figure 5
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    Figure 5

    The structures of three aminoalkylindole cannabinoid receptor agonists: R-(+)-WIN55212, JWH-015, and L-768242.

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    Figure 6

    The structures of five endogenous cannabinoids.

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    Figure 7

    The structures of the CB1-selective synthetic cannabinoid receptor agonists, methanandamide, ACEA, ACPA, and O-1812.

  • Figure 8
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    Figure 8

    The structures of the cannabinoid receptor antagonists/inverse agonists, SR141716A, AM251, AM281, SR144528, and LY320135.

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    Figure 9

    The structures of the pravadoline analogs, AM630, WIN56098, and WIN54461 (6-bromopravadoline).

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    Figure 10

    The structures of O-1184 and O-1238.

  • Figure 11
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    Figure 11

    Amino acid sequence alignment of human, rat, and mouse CB1 and CB2 receptors. Consensus matches are boxed and shaded with darker shading for identities and lighter shading for conservative substitutions. Numbering corresponds to the rat/mouse CB1 sequence. Underlines indicate the positions of the seven transmembrane helices. Helix 3 spans two lines as indicated by the arrowheads on the underline. The rat CB2sequence is a consensus of GenBank accession nos. AF286721 and AF176350together with edited trace data from the rat genome sequencing project (http://www.ncbi.nlm.nih.gov/genome/seq/RnBlast.html). The rat CB2 residue at alignment position 310 appears to be polymorphic [i.e., either Ala (as shown) or Thr].

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    Figure 12

    Autoradiographs show cannabinoid receptor binding (a, f, g) and CB1 (b, d) and CB2 (c, e) mRNA expression in sections from the mouse (sagittal) and human brain (coronal) and mouse spleen (M. Herkenham and A. Hohmann, unpublished observations). Receptor binding of [3H]CP55940, a high-affinity agonist, shows high levels of receptors in the basal ganglia, cerebellum, hippocampus (hipp), and cerebral cortex (a). Cells expressing CB1 mRNA are shown in a similar plane of section (b). Lack of detectable CB2 expression in brain (c) indicates that the binding is to the CB1 type. In contrast, spleen has the opposite relative abundance of CB1 (d) versus CB2 mRNA (e) expression. The human brain has a distribution of cannabinoid receptors that closely matches that of the mouse, with high levels expressed in the basal ganglia, intermediate levels in the amygdala and hypothalamus, and low levels in the thalamus (f, g). The high levels of binding in many areas [cerebellar molecular layer, globus pallidus (GP, GPe), entopeduncular nucleus (Ep, GPi), substantia nigra pars reticulata (SNR), and dentate gyrus molecular layer] are on axons of cells expressing mRNA in afferent areas, such as the caudate putamen (CPu). Some cells in cortex and hippocampus express extremely high levels of CB1 message (arrows in b). Bars measure 1 mm for mouse and 1 cm for human.

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    Figure 13

    Electron micrograph of consecutive rat brain hippocampal sections stained with the C terminus-CB1antibody showing that inhibitory terminals presynaptically express CB1 cannabinoid receptors in the hippocampus. Serial sections have been cut through a CB1-immunoreactive axon terminal forming a symmetrical (GABAergic) synapse (thick arrow) on a dendrite in the stratum radiatum of the CA1 region. Gold particle labeling (small arrows) is restricted to the inner surface of the bouton, where the intracellular carboxy terminus epitope of CB1 is located. A small arrowhead indicates a dense core vesicle. In contrast, the complete lack of staining in axon terminals (★), forming an asymmetrical synapse (large arrowhead), suggests that glutamatergic axons do not contain CB1 receptors. Scale bar is 0.2 μm. Courtesy of I. Katona and T. F. Freund.

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    Figure 14

    The structures of abnormal cannabidiol and O-1602.

Tables

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    Table 1

    Recent reviews on cannabinoid receptors or endogenous cannabinoids

    CoverageAuthors
    Pharmacology, coupling, localization Howlett, 1995a,b;Pertwee, 1997; Felder and Glass, 1998; Ameri, 1999
    Agonists and antagonists Barth and Rinaldi-Carmona, 1999; Pertwee, 1999
    Signal transduction Howlett and Mukhopadhyay, 2000
    Localization and function of CB1 receptors in the central nervous system Elphick and Egertová, 2001
    Molecular biology Onaivi et al., 1996; Matsuda, 1997
    Molecular modeling Reggio, 1999
    Regulation of immune response, coupling Berdyshev, 2000; Cabral, 2001
    Biochemistry and pharmacology of the endocannabinoids Mechoulam et al., 1998; Di Marzo et al., 1999; Martin et al., 1999; Palmer et al., 2000; Reggio and Traore, 2000
    Behavioral effects of cannabinoids in animals Chaperon and Thiebot, 1999
    Cannabinoid receptors and neurotransmitter release Schlicker and Kathman, 2001
    Cannabinoid receptors and pain Martin and Lichtman, 1998; Pertwee, 2001b
    Therapeutic potential Pertwee, 2000b; Piomelli et al., 2000;Porter and Felder, 2001
    • View popup
    Table 2

    Ki values of certain ligands for the in vitro displacement of [3H]CP55940, [3H]R-(+)-WIN55212, or [3H]HU-243 from CB1- and CB2-specific binding sites

    LigandCB1KiValueCB2Ki ValueReference
    nM
    CB1-selective ligands in order of decreasing CB1/CB2 selectivity
     ACEA1.42-a 2-b >2,0002-a 2-b Hillard et al., 1999
     O-18123.42-b 3,8702-b Di Marzo et al., 2001a
     SR141716A11.813,200 Felder et al., 1998
    11.8973 Felder et al., 1995
    12.3702 Showalter et al., 1996
    5.6>1,000 Rinaldi-Carmona et al., 1994
    1.982-b >1,0002-b Rinaldi-Carmona et al., 1994
     AM281122-b 4,2002-c Lan et al., 1999a
     ACPA2.22-a 2-b 7152-a 2-b Hillard et al., 1999
     2-Arachidonylglyceryl ether21.22-b >3,000 Hanus et al., 2001
     LY32013514114,900 Felder et al., 1998
     R-(+)-methanandamide17.92-a 2-b 8682-a 2-c Lin et al., 1998
    202-a 2-b 8152-c Khanolkar et al., 1996
    Ligands without any marked CB1/CB2 selectivity
     Anandamide612-a 2-b 1,9302-a 2-c Lin et al., 1998
    892-a 3712-a Showalter et al., 1996
    5431,940 Felder et al., 1995
    71.72-a 2-b 2792-a 2-b Hillard et al., 1999
    2522-b 581 Mechoulam et al., 1995
     2-Arachidonoylglycerol4722-b 1,400 Mechoulam et al., 1995
    58.32-d 1452-d Ben-Shabat et al., 1998
     HU-2100.06080.524 Felder et al., 1995
    0.12-b 0.17 Rhee et al., 1997
    0.730.22 Showalter et al., 1996
     CP5594051.8 Ross et al., 1999a
    3.722.55 Felder et al., 1995
    1.372-b 1.372-b Rinaldi-Carmona et al., 1994
    0.580.69 Showalter et al., 1996
    0.502-a 2-b 2.802-a 2-b Hillard et al., 1999
     Δ9-THC53.375.3 Felder et al., 1995
    39.52-b 40 Bayewitch et al., 1996
    40.736.4 Showalter et al., 1996
    80.32-b 32.2 Rhee et al., 1997
    35.32-b 3.92-b Rinaldi-Carmona et al., 1994
     Δ8-THC47.62-b 39.32-c Busch-Petersen et al., 1996
     R-(+)-WIN552129.942-b 16.22-b Rinaldi-Carmona et al., 1994
    4.42-a 2-b 1.22-a 2-b Hillard et al., 1999
    1.890.28 Showalter et al., 1996
    62.33.3 Felder et al., 1995
    1234.1 Shire et al., 1996a
    CB2-selective ligands in order of increasing CB2/CB1 selectivity
     JWH-01538313.8 Showalter et al., 1996
     JWH-0511.22-b 0.032 Huffman et al., 1996
     L-7682421,91712 Gallant et al., 1996
     JWH-1392,2902-b 14 Huffman et al., 1998
     AM 6305,15231.2 Ross et al., 1999a
     JWH-1336772-b 3.4 Huffman et al., 1999
     L-7596331,0436.4 Ross et al., 1999a
    15,85020 Gareau et al., 1996
     L-7596564,88811.8 Ross et al., 1999a
    >20,00019.4 Gareau et al., 1996
     HU-308>10,0002-b 22.7 Hanus et al., 1999
     SR1445284370.60 Rinaldi-Carmona et al., 1998
    3052-b 0.302-b Rinaldi-Carmona et al., 1998
    >10,0005.6 Ross et al., 1999a
    • DMH, dimethylheptyl.

    • ↵2-a  With phenylmethylsulfonyl fluoride.

    • ↵2-b  Binding to rat cannabinoid receptors on transfected cells or on brain (CB1) or spleen tissue (CB2).

    • ↵2-c  Binding to mouse spleen cannabinoid receptors.

    • ↵2-d  Species unspecified. All other data from experiments with human cannabinoid receptors.

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    Table 3

    CB1 and CB2 Ki values of stereoisomers of cannabidiol and of two cannabidiol analogs

    LigandCB1Ki ValueCB2Ki ValueReference
    nM
    (−)-CBD4,3502,860 Showalter et al., 1996
    >10,000>10,000 Bisogno et al., 2001
    (+)-CBD842203 Bisogno et al., 2001
    (−)-5′-DMH-CBD>10,0001,800 Bisogno et al., 2001
    (+)-5′-DMH-CBD17.4211 Bisogno et al., 2001
    (−)-7-OH-5′-DMH-CBD4,400671 Bisogno et al., 2001
    (+)-7-OH-5′-DMH-CBD2.544 Bisogno et al., 2001
    • DMH, dimethylheptyl.

    • View popup
    Table 4

    Amino acid sequence variations in CB1 among 60 placental mammals

    Position4-aVariants4-bNumber/TotalConsensusVariants4-bDomain
    539 /59FYAmino terminal
    662 /59DEAmino terminal
    6725 /29NS,T,HAmino terminal
    6822 /59PA,S,TAmino terminal
    694 /59QPAmino terminal
    714 /59VG,I,AAmino terminal
    7312 /59AG,V,SAmino terminal
    744 /59DG,V,AAmino terminal
    7520 /59—P,DAmino terminal
    7620 /59QG,T,A,—Amino terminal
    7715 /58VL,A,T,G,IAmino terminal
    7921 /59IL,M,VAmino terminal
    832 /59YF,LAmino terminal
    9012 /60FYAmino terminal
    9421 /60ED,G,—Amino terminal
    1064 /60MIAmino terminal
    1114 /60IVAmino terminal
    1764 /60VIExtracellular end of TM1
    1875 /60PR,HExtracellular, adjacent to TM3
    2594 /60KEExtracellular, adjacent to TM4
    2626 /60QK,RExtracellular, between TM4 and TM5
    2714 /60LHExtracellular, between TM4 and TM5
    2862 /54TSTM5
    3122 /53RPIntracellular, between TM5 and TM6
    • TM, transmembrane.

    • ↵4-a  Numbering based on rat (or mouse) sequence.

    • ↵4-b  Variant sequences listed only for positions at which more than one sequence deviates from the consensus.

    • View popup
    Table 5

    Detection of cannabinoid receptors in immune cells and tissues

    Cell Type/TissueSpeciesReceptor TypeMethod of Detection
    B lymphocytesHumanCB2 RT-PCR5-aor confocal microscopy5-b
    MacrophagesHuman, mouse, ratCB2 RT-PCR5-a 5-c 5-d
    Mast cellsRatCB2 RT-PCR5-e
    MicrogliaRatCB1, CB2 Mutational RT-PCR,5-d 5-f Western immunoblot,5-d 5-f or immunocytochemistry5-f 5-g
    Natural killer cellsHumanCB2 RT-PCR5-a
    Peripheral mononuclear cellsHuman, ratCB2 RT-PCR5-a 5-e
    CD4 lymphocytesHumanCB2 RT-PCR5-a
    CD8 lymphocytesHumanCB2 RT-PCR5-a
    Lymph nodesHumanCB2 RT-PCR5-a
    Peyer's patchesRatCB5-150 Radioligand binding5-h or radioligand autoradiography5-h
    SpleenHuman, mouse, ratCB1, CB2 Radioligand binding,5-h 5-i radioligand autoradiography,5-h Northern blot,5-j in situ hybridization,5-j or RT-PCR5-a 5-e
    TonsilsHumanCB2 RT-PCR5-aor immunocytochemistry5-a
    ThymusHumanCB2 RT-PCR5-a
    • ↵5-150  CB1 and/or CB2.

    • ↵5-a  Galiègue et al., 1995.

    • ↵5-b  Carayon et al., 1998.

    • ↵5-c  Lee et al., 2001.

    • ↵5-d  Carlisle et al., 2002.

    • ↵5-e  Facci et al., 1995.

    • ↵5-f  Waksman et al., 1999.

    • ↵5-g  Sinha et al., 1998.

    • ↵5-h  Lynn and Herkenham, 1994.

    • ↵5-i  Kaminski et al., 1992.

    • ↵5-j  Munro et al., 1993.

    • View popup
    Table 6

    Cannabinoid-induced inhibition of central and peripheral neurotransmitter release

    TransmitterTissue Preparation or Brain AreaTransmitter-Releasing StimulusReferences
    In vivo
    ACh6-a Rat medial-prefrontal cortexNone Gessa et al., 1998a
    Rat hippocampusNone—6-b
    GABA6-c Rat striatumNone Tersigni and Rosenberg, 1996
    In vitro
    AChRat hippocampal slicesES—6-d
    Rat hippocampal & frontal cortical synaptosomesK+ or Ca2+ Gifford et al., 2000
    Mouse hippocampal or cerebrocortical slicesES or Ca2+ —6-e
    Guinea pig intestinal tissue (MPLM)ES—6-f
    NAGuinea pig cerebrocortical slicesES Schlicker et al., 1997
    Human and guinea pig hippocampal slicesES or Ca2+ Schlicker et al., 1997
    Guinea pig hippocampal slicesNMDA or kainate Kathmann et al., 1999a
    Guinea pig hypothalamic slicesES Schlicker et al., 1997
    Guinea pig cerebellar slicesES Schlicker et al., 1997
    Guinea pig retinal discsES or Ca2+ Schlicker et al., 1996
    Guinea pig bronchial slicesES Vizi et al., 2001
    Human atrial appendage segmentsES Molderings et al., 1999
    Rat atriaES Ishac et al., 1996
    Rat heartES Kurihara et al., 2001
    Rat vas deferensES Ishac et al., 1996
    Mouse vas deferensES Trendelenburg et al., 2000
    Mouse cultured sympathetic neuronsES Göbel et al., 2000
    DARat striatal slicesNMDA Kathmann et al., 1999a
    Rat striatal slicesES Cadogan et al., 1997
    Guinea pig retinal discsES Schlicker et al., 1996
    5-HTMouse cerebrocortical slicesES or Ca2+ Nakazi et al., 2000
    Mouse hypothalamic slicesES Kathmann et al., 1999b
    GABAHuman hippocampal slicesES Katona et al., 2000
    Rat hippocampal slicesES Katona et al., 1999
    GABA6-c Rat or mouse hippocampal slicesES—6-g
    Slices of rat or mouse amygdala (BLC)ES Katona et al., 2001
    1y cultures of neonatal rat hippocampal cellsNone Irving et al., 2000
    1y cultures of neonatal rat hippocampal cellsES Ohno-Shosaku et al., 2001
    Rat striatal slicesES Szabo et al., 1998
    Rat midbrain slices (SNR)ES—6-h
    Rat brain slices (RVM)ES Vaughan et al., 1999
    Rat cerebellar slicesNone Takahashi and Linden, 2000
    Rat cerebellar slicesES Kreitzer and Regehr, 2001b
    Rat brain slices (PAG)ES Vaughan et al., 2000
    Rat brain slices (shell region of NAc)ES Hoffman and Lupica, 2001
    Mouse brain slices (NAc)ES Manzoni and Bockaert, 2001
    Rat spinal trigeminal nucleus pars caudalis (SG)ES Jennings et al., 2001
    Guinea pig intestinal tissue (MPLM)Ethylenediamine Begg et al., 2002
    Glu6-c Rat prefrontal cortical slicesES Auclair et al., 2000
    Rat brain slices (PAG)ES Vaughan et al., 2000
    Mouse brain slices (NAc)ES Robbe et al., 2001
    1y cultures of rat hippocampal cellsLow [Mg2+]o —6-i
    1y cultures of rat hippocampal cellsES Sullivan, 1999
    Mouse hippocampal slices6-j ES—6-k
    Rat or mouse cerebellar slicesES—6-l
    Rat striatal slicesES—6-m
    Rat midbrain slices (SNR)ES Szabo et al., 2000
    Rat spinal cord slices (SG)ES Morisset and Urban, 2001
    1y cultures of rat cerebellar granule cellsLow [Mg2+]o Irving et al., 2001
    Gly6-c Rat spinal trigeminal nucleus pars caudalis (SG)ES Jennings et al., 2001
    d-Asp1y cultures of rat cerebellar granule cellsK+ Breivogel et al., 1999
    CCKRat hippocampal slicesK+ Beinfeld and Connolly, 2001
    • ES, electrical stimulation; [Mg2+]o, extracellular magnesium concentration; MPLM, myenteric plexus-longitudinal muscle preparation; BLC, basolateral complex; NAc, nucleus accumbens; PAG, periaqueductal gray; RVM, rostral ventromedial medulla; SG, substantia gelatinosa; SNR, substantia nigra pars reticulata; ACh, acetylcholine; DA, dopamine; d-Asp,d-aspartate; NA, noradrenaline; 1y, primary.

    • ↵6-a  ACh collected by microdialysis.

    • ↵6-b  Gessa et al., 1997, 1998a; Carta et al., 1998; Nava et al., 2000, 2001.

    • ↵6-c  Indirect electrophysiological evidence for decreased transmitter release: in some of these investigations, there was also evidence that cannabinoids inhibited spontaneous as well as evoked release of GABA or Glu.

    • ↵6-d  Gifford and Ashby, 1996; Gifford et al., 1997a,b; 1999; Kathmann et al., 2001a.

    • ↵6-e  Nakazi et al., 2000; Kathmann et al., 2001a,b.

    • ↵6-f  Pertwee et al., 1996; Coutts and Pertwee, 1997; Mang et al., 2001.

    • ↵6-g  Hájos et al., 2000, 2001;Hoffman and Lupica, 2000; Wilson and Nicoll, 2001.

    • ↵6-h  Chan and Yung, 1998; Chan et al., 1998.

    • ↵6-i  Shen et al., 1996; Shen and Thayer, 1998a,b; 1999.

    • ↵6-j  Signs ofR-(+)-WIN55212-induced inhibition of glutamate release have been observed in tissue from both wild-type and CB1 −/− mice.

    • ↵6-k  Misner and Sullivan, 1999;Hájos et al., 2001.

    • ↵6-l  Lévénès et al., 1998; Kreitzer and Regehr, 2001a; Maejima et al., 2001.

    • ↵6-m  Gerdeman and Lovinger, 2001; Huang et al., 2001.

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Pharmacological Reviews: 54 (2)
Pharmacological Reviews
Vol. 54, Issue 2
1 Jun 2002
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Review ArticleReview

International Union of Pharmacology. XXVII. Classification of Cannabinoid Receptors

A. C. Howlett, F. Barth, T. I. Bonner, G. Cabral, P. Casellas, W. A. Devane, C. C. Felder, M. Herkenham, K. Mackie, B. R. Martin, R. Mechoulam and R. G. Pertwee
Pharmacological Reviews June 1, 2002, 54 (2) 161-202; DOI: https://doi.org/10.1124/pr.54.2.161

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Review ArticleReview

International Union of Pharmacology. XXVII. Classification of Cannabinoid Receptors

A. C. Howlett, F. Barth, T. I. Bonner, G. Cabral, P. Casellas, W. A. Devane, C. C. Felder, M. Herkenham, K. Mackie, B. R. Martin, R. Mechoulam and R. G. Pertwee
Pharmacological Reviews June 1, 2002, 54 (2) 161-202; DOI: https://doi.org/10.1124/pr.54.2.161
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  • Article
    • Abstract
    • I. Introduction: Overview of the Cannabinoid Receptors
    • II. Classification of Ligands That Bind to Cannabinoid Receptors
    • III. Bioassay
    • IV. Cellular Signal Transduction
    • V. Molecular Biology of Cannabinoid Receptors
    • VI. Cannabinoid Receptor Knockout Mice
    • VII. Tissue Distribution of Cannabinoid Receptors
    • VIII. Effects on Neurotransmission
    • IX. Immunological Effects
    • X. Anandamide Is a Vanilloid Receptor Agonist
    • XI. Preliminary Pharmacological Evidence for Non-CB1, Non-CB2 Cannabinoid Receptors
    • XII. Conclusions
    • Footnotes
    • Abbreviations
    • References
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