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Cannabidiol was compared with rimonabant in the brain tissue experiments and with N -[ 1 S -endo-1,3,3-trimethyl bicyclo[2. Accordingly, in some experiments cannabidiol was compared with a putative neutral cannabinoid receptor antagonist, the synthetic cannabidiol analogue, O Thomas et al.
This compound differs from cannabidiol and rimonabant by behaving as a neutral antagonist of cannabinoid receptor agonists rather than as an inverse agonist in the mouse isolated vas deferens Pertwee, In this study, we report first that cannabidiol can behave as an inverse agonist at the human CB 2 receptor. Second, we demonstrate that cannabidiol behaves as a high-potency antagonist of cannabinoid receptor agonists in mouse brain tissue and in membranes from CHO cells transfected with human CB 2 receptors.
Furthermore, the high potency of cannabidiol as an antagonist of the cannabinoid receptor agonist CP in the hCB 2 -CHO cell membranes appears to be a consequence of the ability of cannabidiol to behave as an inverse agonist at the hCB 2 receptor. Some of the results described in this paper have been presented to the International Cannabinoid Research Society Thomas et al.
The methods used comply with the UK Animals Scientific Procedures Act, and associated guidelines for the use of experimental animals. The concentration of [ 3 H]CP used in the displacement assays was 0.
Experiments conducted in native CHO cells were performed under identical conditions to those used for experiments with hCB 2 -CHO cell membranes, whereas the conditions used for the CB 1 -CHO cells were similar to those used for experiments conducted with mouse brain membranes. The vehicle concentration in experiments conducted using one of these drugs was 0.
Values have been expressed as means and variability as s. Its dissociation constant K i was calculated using the equation of Cheng and Prusoff These values were compared with the level of basal binding by performing a one-sample t -test GraphPad Prism. Values for EC 50 , for maximal effect E max and for the s. The apparent dissociation constant K B values for antagonism of agonists by cannabidiol, rimonabant, SR or O have been calculated by Schild analysis from the concentration ratio, defined as the concentration of an agonist that elicits a response of a particular size in the presence of a competitive reversible antagonist at a concentration, B, divided by the concentration of the same agonist that produces an identical response in the absence of the antagonist.
The methods used to determine concentration ratios and apparent K B values and to establish whether log concentration—response plots deviated significantly from parallelism are detailed elsewhere Pertwee et al. Mean values have been compared using Student's two-tailed t- test for unpaired data or one-way analysis of variance ANOVA followed by Bonferroni's multiple comparison test GraphPad Prism.
Both the apparent K B values of rimonabant 0. Mean apparent K B values of these cannabinoids for their antagonism of CP have been calculated from these data and are listed in Table 1. The mean EC 50 and E max values of these cannabinoids are listed in Table 2. Our next experiments were performed with the putative neutral CB 1 receptor antagonist, O Thomas et al. We also found that O shared the ability of cannabidiol to antagonize CP Figure 1c. This finding was unexpected, as Breivogel et al.
We also investigated whether cannabidiol would antagonize ligand-induced activation of a non-cannabinoid G protein-coupled receptor. We selected morphine for these experiments as it is expected to target all the opioid receptor subtypes that are thought to be present in mouse brain membranes Mignat et al. Similar results were obtained from experiments with SR performed under the same assay conditions Figure 7b. Thus, SR induced a downward as well as rightward displacement of the CP log concentration—response curve and exhibited an apparent K B value that was 15 times less than its CB 2 K i value Table 1.
The mean K i values for this displacement were calculated by the Cheng—Prusoff equation and are listed in Table 1. A summary of these results can be found in Table 2. As O attenuated CP responses in mouse brain membranes in a manner that is consistent with it being a neutral CB 1 receptor antagonist, it was of interest to determine whether O also behaves as a neutral antagonist at the CB 2 receptor. O appeared to induce downward as well as rightward displacements of the CP log concentration—response curve in the hCB 2 -CHO cell membranes in these experiments Figure 7c.
However, in contrast to our findings with cannabidiol and SR, this downward displacement was not statistically significant. The results described in this paper indicate that the unexpectedly high potency reported previously for cannabidiol-induced antagonism of cannabinoid agonists in the mouse vas deferens Pertwee et al. However, they are similar to the corresponding apparent K B values We have also found in a previous investigation Pertwee et al.
Thus, the apparent K B values of rimonabant for antagonism of these two cannabinoid receptor agonists were respectively 24 and 7 times lower than the K i of rimonabant for its displacement of [ 3 H]CP from mouse brain membranes. Interestingly, in experiments using assay conditions almost identical to those used in the present investigation, Breivogel et al.
Thus, Savinainen et al. Moreover, cannabidiol has recently been found to inhibit the cellular uptake of adenosine Carrier et al. Moreover the experiments in which Breivogel et al. Similar results were obtained with SR SR has been reported previously to behave as an inverse agonist at the CB 2 receptor Bouaboula et al.
Although, there still appears to be a downward displacement of the CP log concentration—response curve, this was not found to be statistically significant. This hypothesis is supported by results obtained with O In conclusion, this paper provides evidence that cannabidiol exhibits unexpectedly high potency in vitro as an antagonist of both CB 1 and CB 2 receptor agonists and that this antagonism is non-competitive in nature.
It is noteworthy, however, that Breivogel et al. As to the high potency displayed by cannabidiol as an antagonist of CB 2 receptor activation, our data suggest that this may stem from its ability to induce CB 2 receptor inverse agonism at concentrations well below those at which it displaces [ 3 H]CP from these receptors. This action may also contribute to the well-known anti-inflammatory properties of cannabidiol reviewed in Pertwee, , as there is evidence that CB 2 receptor inverse agonism can inhibit immune cell migration Lunn et al.
O may also be a neutral CB 2 receptor antagonist.
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