The use of cannabinoids in animals and therapeutic implications for veterinary medicine: a review

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Veterinarni Medicina, 61, 2016 (3): 111–122 doi: 10.17221/8762-VETMED 2001), O-arachidonoylethanolamine (virhodamine) (Porter et al. 2002) and N-arachidonoyldopamine (NADA) (Bisogno et al. 2000; Gaffuri et al. 2012; Mechoulam et al. 2014). Within the nervous system endocannabinoids are released from post-synaptic neurons (retrograde neurotransmission) and they bind to presynaptic CB1 receptors (see below) which results particularly in inhibition of GABA or glutamate release (Heifets and Castillo 2009). In neuron-astrocyte signalling cannabinoids released from post-synaptic neurons stimulate astrocytic CB1 receptors, thereby triggering glutamatergic gliotransmission (Castillo et al. 2012). Phytocannabinoids are chemicals produced es- pecially by female plants of Cannabis sativa and are present in the resin of the herb. It has been found that these plants contain over 100 phyto- cannabinoids (Hill et al. 2012). The most studied cannabinoids from Cannabis sativa include e.g. delta-9-tetrahydrocannabinol (THC), cannabidiol, tetrahydrocannabivarin, tetrahydrocannabiorcol, cannabichromene and cannabigerol (Maione et al. 2013). THC was first isolated in 1964 (Gaoni and Mechoulam 1964) and the majority of the herbal cannabinoids soon after. Synthetic cannabinoids are manufactured com- pounds which bind to cannabinoid receptors (with either agonistic or antagonistic activity) and many of them were originally synthesised for research purposes in University scientific departments or pharmaceutical companies. The most frequently reported series are represented by JWH (John W. Huffman, Clemson University), CP (Pfizer), HU (Hebrew University), AM (Alexandros Makriyannis, Northeastern University), WIN (Sterling Winthrop) and RCS (Research Chemical Supply) (Presley et al. 2013). Both phytocannabinoids and synthetic can- nabinoids mimic the effects of endocannabinoids (Grotenhermen 2006). Two cannabinoid receptors were initially recog- nised, CB1 and CB2. Both these subtypes belong to the large family of receptors that are coupled to G proteins (Svizenska et al. 2008). Cannabinoid CB1 receptors are among the most plentiful and widely distributed receptors coupled to G proteins in the brain (Grotenhermen 2006). The CB1 receptor was cloned in 1990 (Matsuda et al. 1990) and CB2 in 1993 (Munro et al. 1993). CB1 receptors are present primarily in the central nervous system in regions of the brain that are responsible for pain modula- tion (certain parts of the spinal cord, periaqueduct- Review Article al grey), movement (basal ganglia, cerebellum) or memory processing (hippocampus, cerebral cortex) (Grotenhermen 2006). To a lesser extent, they can also be found in some peripheral tissues such as pituitary gland, immune cells, reproductive tissues, gastrointestinal tissues, sympathetic ganglia, heart, lung, urinary bladder and adrenal gland (Pertwee 1997). CB2 receptors are particularly expressed in the periphery, in the highest density on immune cells, especially B-cells and natural killer cells (Pertwee 1997) and also in tonsils or spleen (Galiegue et al. 1995); nevertheless, their presence has also been described in the CNS (Van Sickle et al. 2005). The frequently discussed psychotropic effects of can- nabinoids are mediated only by the activation of CB1 receptors and not of CB2 receptors (Grotenhermen and Muller-Vahl 2012). Endocannabinoids have also been shown to act on TRPV1 receptors (transient receptor potential cat- ion channels subfamily V member 1, also known as the “capsaicin receptor” and “vanilloid receptor” 1) (Ross 2003). The existence of other G-protein can- nabinoid receptors has also been suggested. These proposed receptors (also called putative or non- classical cannabinoid receptors) include GPR18, GPR55 and GPR119 that have structural similarity to CB1 and CB2 (Alexander et al. 2013; Zubrzycki et al. 2014). 3. The use of cannabinoids in animals It has been shown that the mechanism of action of cannabinoids is very complex. The activation of cannabinoid CB1 receptors results in retrograde inhibition of the neuronal release of acetylcholine, dopamine, GABA, histamine, serotonin, glutamate, cholecystokinin, D-aspartate, glycine and no- radrenaline (Grotenhermen and Muller-Vahl 2012). CB2 receptors localised mainly in cells associated with the immune system are involved in the control of inflammatory processes. Their activation results in, among other effects, inhibition of pro-inflam- matory cytokine production and increased release of anti-inflammatory cytokines (Zubrzycki et al. 2014). In addition, some cannabinoids were shown to act not only at cannabinoid receptors but also at vanilloid or serotonin 5-HT3 receptors (Contassot et al. 2004; Grotenhermen and Muller-Vahl 2012). This complexity of interactions explains both the 113

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