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{"title"=>"The orexigenic effect of ghrelin is mediated through central activation of the endogenous cannabinoid system", "type"=>"journal", "authors"=>[{"first_name"=>"Blerina", "last_name"=>"Kola", "scopus_author_id"=>"6602919169"}, {"first_name"=>"Imre", "last_name"=>"Farkas", "scopus_author_id"=>"7007056966"}, {"first_name"=>"Mirjam", "last_name"=>"Christ-Crain", "scopus_author_id"=>"6601936720"}, {"first_name"=>"Gábor", "last_name"=>"Wittmann", "scopus_author_id"=>"7007014583"}, {"first_name"=>"Francesca", "last_name"=>"Lolli", "scopus_author_id"=>"24344127100"}, {"first_name"=>"Faisal", "last_name"=>"Amin", "scopus_author_id"=>"24449180300"}, {"first_name"=>"Judith", "last_name"=>"Harvey-White", "scopus_author_id"=>"6701653318"}, {"first_name"=>"Zsolt", "last_name"=>"Liposits", "scopus_author_id"=>"7004411568"}, {"first_name"=>"George", "last_name"=>"Kunos", "scopus_author_id"=>"7006126817"}, {"first_name"=>"Ashley B.", "last_name"=>"Grossman", "scopus_author_id"=>"35401342800"}, {"first_name"=>"Csaba", "last_name"=>"Fekete", "scopus_author_id"=>"7006236470"}, {"first_name"=>"Márta", "last_name"=>"Korbonits", "scopus_author_id"=>"7004190977"}], "year"=>2008, "source"=>"PLoS ONE", "identifiers"=>{"scopus"=>"2-s2.0-45549092924", "sgr"=>"45549092924", "issn"=>"19326203", "doi"=>"10.1371/journal.pone.0001797", "pmid"=>"18335063", "isbn"=>"1932-6203 (Electronic)", "pui"=>"351917350"}, "id"=>"11d50474-11da-3734-b9cb-4e856aae02d6", "abstract"=>"INTRODUCTION: Ghrelin and cannabinoids stimulate appetite, this effect possibly being mediated by the activation of hypothalamic AMP-activated protein kinase (AMPK), a key enzyme in appetite and metabolism regulation. The cannabinoid receptor type 1 (CB1) antagonist rimonabant can block the orexigenic effect of ghrelin. In this study, we have elucidated the mechanism of the putative ghrelin-cannabinoid interaction.\\n\\nMETHODS: The effects of ghrelin and CB1 antagonist rimonabant in wild-type mice, and the effect of ghrelin in CB1-knockout animals, were studied on food intake, hypothalamic AMPK activity and endogenous cannabinoid content. In patch-clamp electrophysiology experiments the effect of ghrelin was assessed on the synaptic inputs in parvocellular neurons of the hypothalamic paraventricular nucleus, with or without the pre-administration of a CB1 antagonist or of cannabinoid synthesis inhibitors.\\n\\nRESULTS AND CONCLUSIONS: Ghrelin did not induce an orexigenic effect in CB1-knockout mice. Correspondingly, both the genetic lack of CB1 and the pharmacological blockade of CB1 inhibited the effect of ghrelin on AMPK activity. Ghrelin increased the endocannabinoid content of the hypothalamus in wild-type mice and this effect was abolished by rimonabant pre-treatment, while no effect was observed in CB1-KO animals. Electrophysiology studies showed that ghrelin can inhibit the excitatory inputs on the parvocellular neurons of the paraventricular nucleus, and that this effect is abolished by administration of a CB1 antagonist or an inhibitor of the DAG lipase, the enzyme responsible for 2-AG synthesis. The effect is also lost in the presence of BAPTA, an intracellular calcium chelator, which inhibits endocannabinoid synthesis in the recorded parvocellular neuron and therefore blocks the retrograde signaling exerted by endocannabinoids. In summary, an intact cannabinoid signaling pathway is necessary for the stimulatory effects of ghrelin on AMPK activity and food intake, and for the inhibitory effect of ghrelin on paraventricular neurons.", "link"=>"http://www.mendeley.com/research/orexigenic-effect-ghrelin-mediated-through-central-activation-endogenous-cannabinoid-system", "reader_count"=>112, "reader_count_by_academic_status"=>{"Unspecified"=>2, "Professor > Associate Professor"=>9, "Student > Doctoral Student"=>7, "Researcher"=>29, "Student > Ph. D. Student"=>18, "Student > Postgraduate"=>7, "Student > Master"=>13, "Other"=>5, "Student > Bachelor"=>15, "Lecturer"=>2, "Professor"=>5}, "reader_count_by_user_role"=>{"Unspecified"=>2, "Professor > Associate Professor"=>9, "Student > Doctoral Student"=>7, "Researcher"=>29, "Student > Ph. D. Student"=>18, "Student > Postgraduate"=>7, "Student > Master"=>13, "Other"=>5, "Student > Bachelor"=>15, "Lecturer"=>2, "Professor"=>5}, "reader_count_by_subject_area"=>{"Unspecified"=>7, "Biochemistry, Genetics and Molecular Biology"=>3, "Nursing and Health Professions"=>1, "Medicine and Dentistry"=>19, "Agricultural and Biological Sciences"=>53, "Neuroscience"=>18, "Design"=>1, "Pharmacology, Toxicology and Pharmaceutical Science"=>5, "Psychology"=>2, "Chemistry"=>2, "Social Sciences"=>1}, "reader_count_by_subdiscipline"=>{"Design"=>{"Design"=>1}, "Medicine and Dentistry"=>{"Medicine and Dentistry"=>19}, "Neuroscience"=>{"Neuroscience"=>18}, "Chemistry"=>{"Chemistry"=>2}, "Social Sciences"=>{"Social Sciences"=>1}, "Psychology"=>{"Psychology"=>2}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>53}, "Nursing and Health Professions"=>{"Nursing and Health Professions"=>1}, "Biochemistry, Genetics and Molecular Biology"=>{"Biochemistry, Genetics and Molecular Biology"=>3}, "Unspecified"=>{"Unspecified"=>7}, "Pharmacology, Toxicology and Pharmaceutical Science"=>{"Pharmacology, Toxicology and Pharmaceutical Science"=>5}}, "reader_count_by_country"=>{"Netherlands"=>1, "United States"=>3, "Brazil"=>1, "United Kingdom"=>1, "Kenya"=>1, "Portugal"=>1, "Germany"=>1, "Croatia"=>1, "Spain"=>2}, "group_count"=>6}

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• {"type"=>"COMMENT", "annotationUri"=>"info:doi/10.1371/annotation/b20f751a-c0d5-46b2-9827-e25272e75001", "title"=>"Referee comments: Referee 1 (Vincenzo Di Marzo)", "body"=>"Referee 1's review (Vincenzo Di Marzo):\n\nReview of the original manuscript:\nIn this article, Kola et al address the very important issue of whether or not the actions of ghrelin on food intake, AMPK activation and PVN anorexic neuron activity are directly mediated by CB1 receptors and endocannabinoids. The data presented strongly argue in favour of such a possibility. The study has been mostly well performed and the article is well written. The data are very important since they fill a crucial gap in our knowledge of the mechanism of action of both ghrelin and endocannabinoids. I have, however, some major points that need to be addressed by the authors and should strengthen this important study: \n1) the hypothalamic levels of 2-AG seem to be considerably (almost 10-fold) higher than those previously reported by others in the field. Furthermore, it is not clear why, whilst rimonabant reduces 2-AG levels, these levels are higher in CB1 KO mice. In a previous study, surprisingly not quoted by the authors, Di Marzo et al (J Neurochem, 2000) showed no changes in either 2-AG or anandamide levels in the hypothalamus of both male and female CB1 KO mice seemingly of the same strain, background and age as those used in this study. Furthermore, the authors refer to a preliminary work (ref. 20) by Bequet et al., which has never been published in full. These authors, however, did find that rimonabant causes changes in hypothalamic endocannabinoid levels, but the changes found with anandamide were opposite to those with 2-AG, whereas the present authors report here similar changes for 2-AG and anandamide. All these discrepancies should be discussed rather than just being ignored, and, importantly, the results with anandamide should be also shown. \n2) The authors have reported effects of rimonabant on ghrelin-induced AMPK and endocannabinoid levels. Yet, they seem to have used a per se non-inactive dose of rimonabant (Fig. 1b,c). It is possible that what they see is not antagonism, but rather the sum of two opposite effects. Control experiments with an inactive per se dose of rimonabant should be carried out; \n3) Given for granted that the authors deal with the above two points, they should discuss further why the effect of ghrelin on endocannabinoid levels is not observed in the presence of inactive CB1 receptors. They mention a positive feed-back loop on endocannabinoid biosynthesis. This possibility might be supported by the previous finding that also NPY is inactive in CB1 KO mice (Poncelet et al., Neurosci Lett. 2003) and yet that CB1 stimulation enhances NPY levels (Gamber et al., Neuropharmacology, 2005), two other very relevant papers that should have been quoted by the authors. It is possible that ghrelin stimulates EC levels both directly and indirectly via NPY, whose levels are then further amplified by CB1 stimulation, thus leading to further EC biosynthesis. This positive feed-back mechanism would require both ghrelin and CB1 receptors. By converse, tonic CB1 receptor stimulation alone (without concomitant ghrelin receptor stimulation, necessary to trigger both NPY and endocannabinoid levels) might not be sufficient to elevate EC levels, and this might explain why CB1 KO mice do not exhibit lower EC levels in the hypothalamus. The authors might wish to discuss this possibility. \n4) The electrophysiology experiments are very important but perhaps should be accompanied by a little bit more pharmacology. For example, in order to establish a true link between the effects of ghrelin on food-intake and EC levels and its effect on PVN neuron activity, it would be important to see if a DAGLalpha inhibitor also blocks the effect of ghrelin in these experiments. In fact, rimonabant might behave as an inverse agonist in vitro, and BAPTA-AM is not specific for endocannabinoid biosynthesis. DAGL inhibitors such as THC have been widely used recently in similar studies on the role of endocannabinoids as retrograde messengers. \n\nMinor point: 1) since the effect of ghrelin on endocannabinoid levels is a crucial part of this work, recent reviews and experimental papers showing the effects, or lack thereof, of other hypothalamic hormones on endocannabinoid levels should be quoted in the references. 2) The name of the last author in reference 9 is mispelt.\n\nReview of the first revised manuscript:\nThe authors have answered satisfactorily to my comments\n\n**********\nN.B. These are the comments made by the referee when reviewing an earlier version of this paper. Prior to publication the manuscript has been revised in light of these comments and to address other editorial requirements.", "isRemoved"=>false, "created"=>"2008-03-18T10:45:09Z", "lastModified"=>"2008-03-18T10:45:09Z", "creator"=>{"userId"=>"80539"}, "highlightedText"=>"", "competingInterestStatement"=>{"creatorWasPrompted"=>false, "hasCompetingInterests"=>false}, "parentArticle"=>{"doi"=>"info:doi/10.1371/journal.pone.0001797", "state"=>"published", "journals"=>{"PLoSONE"=>{"journalKey"=>"PLoSONE", "eIssn"=>"1932-6203", "title"=>"PLOS ONE"}}}, "replyTreeSize"=>0, "mostRecentActivity"=>"2008-03-18T10:45:09Z", "replies"=>[]}
• {"type"=>"COMMENT", "annotationUri"=>"info:doi/10.1371/annotation/4cba1ad9-1379-47ce-b46d-e1beabdc0e7e", "title"=>"Referee comments: Referee 2", "body"=>"Referee 2's review:\n\nThe authors revised their manuscript and provide new sets of data. Overall, relatively solid circumstantial evidence is provided to suggest that ghrelin's action is mediated by CB1 receptors.\n\nHowever, the data is not conclusive and remains to be indirect. Specifically, the reluctance of the authors to provide thorough electrophysiological proof that ghrelin is not capable generating increased or decreased action potentials in any neuronal system of CB1 KO mice is unfortunate. There are numerous publications by various laboratories analyzing ghrelin's action on neuronal firing in various hypothalamic and extrahypothalamic sites without GFP guidance. The paper by Luquet et al. (Luquet S, Phillips CT, Palmiter RD. NPY/AgRP neurons are not essential for feeding responses to glucoprivation.Peptides. 2007 Feb;28(2):214-25. Epub 2006 Dec 27.) clearly shows the pivotal role of NPY/AgRP neurons in mediating ghrelin's effect on feeding. The PVN, which may be affected by ghrelin, cannot be a surrogate for such a critical conclusion to be made. NPY/AgRP neurons have been recorder from without a GFP construct in rats: van den Top M, Lyons DJ, Lee K, Coderre E, Renaud LP, Spanswick D. Pharmacological and molecular characterization of ATP-sensitive K(+) conductances\nin CART and NPY/AgRP expressing neurons of the hypothalamic arcuate nucleus. Neuroscience. 2007 Feb 9;144(3):815-24. Epub 2006 Nov 28.; van den Top M, Lee K, Whyment AD, Blanks AM, Spanswick D. Orexigen-sensitive NPY/AgRP pacemaker neurons in the hypothalamic arcuate nucleus. Nat Neurosci. 2004 May;7(5):493-4. Epub 2004 Apr 18. Is it difficult to carry out these experiments? Yes. But are there necessary to reach a reliable conclusion? In the view of the reviewers is yes.\n\n**********\nN.B. These are the comments made by the referee when reviewing an earlier version of this paper. Prior to publication the manuscript has been revised in light of these comments and to address other editorial requirements.", "isRemoved"=>false, "created"=>"2008-03-18T10:47:22Z", "lastModified"=>"2008-03-18T10:47:22Z", "creator"=>{"userId"=>"80539"}, "highlightedText"=>"", "competingInterestStatement"=>{"creatorWasPrompted"=>false, "hasCompetingInterests"=>false}, "parentArticle"=>{"doi"=>"info:doi/10.1371/journal.pone.0001797", "state"=>"published", "journals"=>{"PLoSONE"=>{"journalKey"=>"PLoSONE", "eIssn"=>"1932-6203", "title"=>"PLOS ONE"}}}, "replyTreeSize"=>0, "mostRecentActivity"=>"2008-03-18T10:47:22Z", "replies"=>[]}

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• {"id"=>"910706037250039808", "text"=>"#AMPK #CB@1 The Orexigenic Effect of Ghrelin Is Mediated through Central Activation of the Endogenous Cannabino ... https://t.co/LGHOj4zZhs", "created_at"=>"2017-09-21T03:23:34Z", "user"=>"Oxyconmdamu", "user_name"=>"Bearzapine,PharmC", "user_profile_image"=>"http://pbs.twimg.com/profile_images/868633428039696384/r6y1mReb_normal.jpg"}

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Figshare

• {"files"=>["https://ndownloader.figshare.com/files/938026"], "description"=>"<p>Application of ghrelin (100 nM) in the extracellular solution decreased the amplitude and frequency of the mEPSCs (<a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0001797#pone-0001797-g002\" target=\"_blank\">Fig. 2A–C</a>). Extracellular administration of the cannabinoid receptor antagonist AM251 (1 µM), however, blocked the effect of ghrelin. Both amplitude and frequency changes were attenuated (<a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0001797#pone-0001797-g002\" target=\"_blank\">Fig. 2D–F</a>). In addition, intracellularly applied BAPTA, an effective chelator of free calcium, (<a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0001797#pone-0001797-g002\" target=\"_blank\">Fig. 2G–I</a>) and the DAG lipase (DAGL) inhibitor THL (5 µM) (<a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0001797#pone-0001797-g002\" target=\"_blank\">Fig. 2J–L</a>) also abolished the effect of ghrelin.</p>", "links"=>[], "tags"=>["clamp", "recordings", "mepscs", "parvocellular", "neurons"], "article_id"=>608464, "categories"=>["Computational Biology", "Biochemistry", "Medicine"], "users"=>["Blerina Kola", "Imre Farkas", "Mirjam Christ-Crain", "Gábor Wittmann", "Francesca Lolli", "Faisal Amin", "Judith Harvey-White", "Zsolt Liposits", "George Kunos", "Ashley B. Grossman", "Csaba Fekete", "Márta Korbonits"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0001797.g002", "stats"=>{"downloads"=>3, "page_views"=>139, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Whole_cell_patch_clamp_recordings_of_mEPSCs_in_the_parvocellular_neurons_of_the_PVN_/608464", "title"=>"Whole-cell patch clamp recordings of mEPSCs in the parvocellular neurons of the PVN.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 11:07:16"}
• {"files"=>["https://ndownloader.figshare.com/files/938253"], "description"=>"<p>Binding of ghrelin to its receptor (growth hormone secretagogue receptor 1a, GHS-R1a) on the surface of parvocellular neurons results in an increase of intracellular Ca²⁺ levels due to mobilization of Ca²⁺ from intracellular stores and opening Ca²⁺ channels. The increased intracellular Ca²⁺ level activates the 2-AG synthesizing enzyme diacylglycerol lipase-α (DAGL), directly or through activation of protein kinase C (PKC), resulting in an increased 2-AG synthesis and release into the extracellular space. The increased activation of presynaptic CB1 then inhibits the release of the excitatory neurotransmitter glutamate (Glu) from the axons innervating the PVN neurons. Intracellular administration of the Ca²⁺-chelator BAPTA blocks this cascade by preventing the increase of intracellular Ca²⁺ level, whereas extracellularly-given THL blocks the 2-AG synthesis and AM251 blocks the cascade at the level of CB1.</p>", "links"=>[], "tags"=>["illustrates", "parvocellular", "neurons"], "article_id"=>608691, "categories"=>["Computational Biology", "Biochemistry", "Medicine"], "users"=>["Blerina Kola", "Imre Farkas", "Mirjam Christ-Crain", "Gábor Wittmann", "Francesca Lolli", "Faisal Amin", "Judith Harvey-White", "Zsolt Liposits", "George Kunos", "Ashley B. Grossman", "Csaba Fekete", "Márta Korbonits"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0001797.g004", "stats"=>{"downloads"=>3, "page_views"=>24, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Schematic_drawing_illustrates_the_proposed_model_of_ghrelin_s_action_in_the_parvocellular_neurons_of_the_PVN_/608691", "title"=>"Schematic drawing illustrates the proposed model of ghrelin's action in the parvocellular neurons of the PVN.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 11:08:44"}
• {"files"=>["https://ndownloader.figshare.com/files/937841"], "description"=>"<p>(A) Cumulative food intake of WT and CB1-KO mice after 2 hours of treatment with icv ghrelin (G) or vehicle (C), n = 5–9 mice/group. (B) Ghrelin and cannabinoid effects on hypothalamic AMPK activity one hour after intraperitoneal administration of THC, ghrelin, rimonabant (R) or a combination of rimonabant and ghrelin (R+G) in WT and CB1-KO animals compared to vehicle-treated animals, n = 6–14 mice/group. (C) Hypothalamic 2-AG content after intraperitoneal treatment with ghrelin in WT and CB1-KO animals, n = 6 mice/group. (D) Hypothalamic 2-AG content in WT animals after ip treatment with ghrelin and rimonabant compared to vehicle treated animals, n = 6–14 mice/group. (E) Hypothalamic AEA content after ip treatment with ghrelin in WT and CB1-KO animals. All data shown as mean±SEM, *P<0.05, **P<0.01, ***P<0.001, ⁺P = 0.052.</p>", "links"=>[], "tags"=>["ghrelin", "cannabinoids", "hypothalamic", "ampk", "endocannabinoid"], "article_id"=>608276, "categories"=>["Computational Biology", "Biochemistry", "Medicine"], "users"=>["Blerina Kola", "Imre Farkas", "Mirjam Christ-Crain", "Gábor Wittmann", "Francesca Lolli", "Faisal Amin", "Judith Harvey-White", "Zsolt Liposits", "George Kunos", "Ashley B. Grossman", "Csaba Fekete", "Márta Korbonits"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0001797.g001", "stats"=>{"downloads"=>0, "page_views"=>7, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_effect_of_ghrelin_and_cannabinoids_on_food_intake_hypothalamic_AMPK_activity_and_endocannabinoid_content_/608276", "title"=>"The effect of ghrelin and cannabinoids on food intake, hypothalamic AMPK activity and endocannabinoid content.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 11:06:06"}
• {"files"=>["https://ndownloader.figshare.com/files/938145"], "description"=>"<p>Application of ghrelin resulted in significant changes in all of these parameters whereas administration of AM251, BAPTA or THL eliminated the changes (<a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0001797#pone-0001797-g003\" target=\"_blank\">Fig. 3A–D</a>). All data shown as mean±SEM, α and β correspond to P<0.05 and P<0.01, respectively, and refer to the comparison of ghrelin vs. control. *P<0.05 and **P<0.01 correspond to the comparison between ghrelin vs. AM251+ghrelin, ghrelin vs. BAPTA+ghrelin or ghrelin vs. THL+ghrelin groups.</p>", "links"=>[], "tags"=>["mepscs", "instantaneous", "interevent", "elicited", "ghrelin", "am251", "bapta", "thl", "parvocellular"], "article_id"=>608575, "categories"=>["Computational Biology", "Biochemistry", "Medicine"], "users"=>["Blerina Kola", "Imre Farkas", "Mirjam Christ-Crain", "Gábor Wittmann", "Francesca Lolli", "Faisal Amin", "Judith Harvey-White", "Zsolt Liposits", "George Kunos", "Ashley B. Grossman", "Csaba Fekete", "Márta Korbonits"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0001797.g003", "stats"=>{"downloads"=>3, "page_views"=>43, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Changes_of_various_parameters_of_mEPSCs_A_amplitude_B_instantaneous_frequency_C_interevent_interval_D_event_frequency_elicited_by_ghrelin_as_percentage_of_control_by_AM251_ghrelin_as_percentage_of_AM251_alone_BAPTA_ghrelin_as_percentage_of_BAPTA_alone_an/608575", "title"=>"Changes of various parameters of mEPSCs [(A) amplitude, (B) instantaneous frequency, (C) interevent interval, (D) event frequency] elicited by ghrelin (as percentage of control), by AM251+ghrelin (as percentage of AM251 alone), BAPTA+ghrelin (as percentage of BAPTA alone) and THL+ghrelin (as percentage of THL alone) in the parvocellular neurons.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-02-21 11:08:01"}

PMC Usage Stats | Further Information

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Relative Metric

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F1000Prime