{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T20:58:22Z","timestamp":1788987502934,"version":"build-2803163510"},"reference-count":86,"publisher":"Wiley","issue":"5","license":[{"start":{"date-parts":[[2021,7,15]],"date-time":"2021-07-15T00:00:00Z","timestamp":1626307200000},"content-version":"am","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"},{"start":{"date-parts":[[2021,7,15]],"date-time":"2021-07-15T00:00:00Z","timestamp":1626307200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"DOI":"10.13039\/100011073","name":"Nutrition Obesity Research Center, University of North Carolina","doi-asserted-by":"publisher","award":["P30 DK035816"],"award-info":[{"award-number":["P30 DK035816"]}],"id":[{"id":"10.13039\/100011073","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R01AG061558"],"award-info":[{"award-number":["R01AG061558"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R01CA239208"],"award-info":[{"award-number":["R01CA239208"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000090","name":"Congressionally Directed Medical Research Programs","doi-asserted-by":"publisher","award":["PC170059"],"award-info":[{"award-number":["PC170059"]}],"id":[{"id":"10.13039\/100000090","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000738","name":"U.S. Department of Veterans Affairs","doi-asserted-by":"publisher","award":["BX002807"],"award-info":[{"award-number":["BX002807"]}],"id":[{"id":"10.13039\/100000738","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["J cachexia sarcopenia muscle"],"published-print":{"date-parts":[[2021,10]]},"abstract":"Abstract<\/jats:title>\n \n Background<\/jats:title>\n \n Ghrelin may ameliorate cancer cachexia (CC) by preventing anorexia, muscle, and fat loss. However, the mechanisms mediating these effects are not fully understood. This study characterizes the pathways involved in muscle mass and strength loss in the Lewis lung carcinoma (LLC)\u2010induced cachexia model, and the effects of ghrelin in mice with or without its only known receptor: the growth hormone secretagogue receptor\u20101a ((GHSR\u20101a),\n Ghsr<\/jats:italic>\n \n +\/+<\/jats:italic>\n <\/jats:sup>\n and\n Ghsr<\/jats:italic>\n \n \u2212\/\u2212<\/jats:italic>\n <\/jats:sup>\n ).\n <\/jats:p>\n <\/jats:sec>\n \n Methods<\/jats:title>\n \n Five to 7\u2010month\u2010old male C57BL\/6J\n Ghsr<\/jats:italic>\n \n +\/+<\/jats:italic>\n <\/jats:sup>\n and\n Ghsr<\/jats:italic>\n \n \u2212\/\u2212<\/jats:italic>\n <\/jats:sup>\n mice were inoculated with 1\u00a0\u00d7\u00a010\n 6<\/jats:sup>\n heat\u2010killed (HK) or live LLC cells (tumour implantation, TI). When tumours were palpable (7\u00a0days after TI), tumour\u2010bearing mice were injected with vehicle (T\u00a0+\u00a0V) or ghrelin twice\/day for 14\u00a0days (T\u00a0+\u00a0G, 0.8\u00a0mg\/kg), while HK\u2010treated mice were given vehicle (HK\u00a0+\u00a0V). Body weight and grip strength were evaluated before TI and at termination (21\u00a0days after TI). Hindlimb muscles were collected for analysis.\n <\/jats:p>\n <\/jats:sec>\n \n Results<\/jats:title>\n \n Less pronounced body weight (BW) loss (87.70\u00a0\u00b1\u00a00.98% vs. 83.92\u00a0\u00b1\u00a01.23%, percentage of baseline BW in tumour\u2010bearing\n Ghsr<\/jats:italic>\n \n +\/+<\/jats:italic>\n <\/jats:sup>\n vs.\n Ghsr<\/jats:italic>\n \n \u2212\/\u2212<\/jats:italic>\n <\/jats:sup>\n ,\n P<\/jats:italic>\n \u00a0=\u00a00.008), and lower upregulation of ubiquitin\u2010proteasome system (UPS, MuRF1\/\n Trim63<\/jats:italic>\n , 5.71\u00a0\u00b1\u00a01.53\u2010fold vs. 9.22\u00a0\u00b1\u00a01.94\u2010fold\u2010change from\n Ghsr<\/jats:italic>\n \n +\/+<\/jats:italic>\n <\/jats:sup>\n HK\u00a0+\u00a0V in tumour\u2010bearing\n Ghsr<\/jats:italic>\n \n +\/+<\/jats:italic>\n <\/jats:sup>\n vs.\n Ghsr<\/jats:italic>\n \n \u2010\/\u2010<\/jats:italic>\n <\/jats:sup>\n ,\n P<\/jats:italic>\n \u00a0=\u00a00.036) and autophagy markers (\n Becn1<\/jats:italic>\n ,\n Atg5<\/jats:italic>\n ,\n Atg7<\/jats:italic>\n , tumour\u2010bearing\n Ghsr<\/jats:italic>\n \n +\/+<\/jats:italic>\n <\/jats:sup>\n \u00a0<\u00a0\n Ghsr<\/jats:italic>\n \n \u2212\/\u2212<\/jats:italic>\n <\/jats:sup>\n , all\n P<\/jats:italic>\n \u00a0<\u00a00.02) were found in T\u00a0+\u00a0V\n Ghsr<\/jats:italic>\n \n +\/+<\/jats:italic>\n <\/jats:sup>\n vs.\n Ghsr<\/jats:italic>\n \n \u2212\/\u2212<\/jats:italic>\n <\/jats:sup>\n mice. Ghrelin attenuated LLC\u2010induced UPS marker upregulation in both genotypes, [\n Trim63<\/jats:italic>\n was decreased from 5.71\u00a0\u00b1\u00a01.53\u2010fold to 1.96\u00a0\u00b1\u00a00.47\u2010fold in\n Ghsr<\/jats:italic>\n \n +\/+<\/jats:italic>\n <\/jats:sup>\n (T\u00a0+\u00a0V vs. T\u00a0+\u00a0G:\n P<\/jats:italic>\n \u00a0=\u00a00.032) and 9.22\u00a0\u00b1\u00a01.94\u2010fold to 4.72\u00a0\u00b1\u00a01.06\u2010fold in\n Ghsr<\/jats:italic>\n \n \u2212\/\u2212<\/jats:italic>\n <\/jats:sup>\n (T\u00a0+\u00a0V vs. T\u00a0+\u00a0G:\n P<\/jats:italic>\n \u00a0=\u00a00.008)]. Only in\n Ghsr<\/jats:italic>\n \n +\/+<\/jats:italic>\n <\/jats:sup>\n mice ghrelin ameliorated LLC\u2010induced grip strength loss [improved from 89.24\u00a0\u00b1\u00a03.48% to 97.80\u00a0\u00b1\u00a02.31% of baseline (T\u00a0+\u00a0V vs. T\u00a0+\u00a0G:\n P<\/jats:italic>\n \u00a0=\u00a00.042)], mitophagy markers [\n Bnip3<\/jats:italic>\n was decreased from 2.28\u00a0\u00b1\u00a00.56 to 1.38\u00a0\u00b1\u00a00.14\u2010fold (T\u00a0+\u00a0V vs. T\u00a0+\u00a0G:\n P<\/jats:italic>\n \u00a0\u2264\u00a00.05)], and impaired mitochondrial respiration [State 3u improved from 698.23\u00a0\u00b1\u00a073.96 to 934.37\u00a0\u00b1\u00a095.21\u00a0pmol\/min (T\u00a0+\u00a0V vs. T\u00a0+\u00a0G:\n P<\/jats:italic>\n \u00a0\u2264\u00a00.05)], whereas these markers were not improved by ghrelin\n Ghsr<\/jats:italic>\n \n \u2212\/\u2212<\/jats:italic>\n <\/jats:sup>\n . Compared with\n Ghsr<\/jats:italic>\n \n +\/+<\/jats:italic>\n <\/jats:sup>\n ,\n Ghsr<\/jats:italic>\n \n \u2212\/\u2212<\/jats:italic>\n <\/jats:sup>\n tumour\u2010bearing mice also showed decreased response to ghrelin in BW [T\u00a0+\u00a0G\u2010treated\n Ghsr<\/jats:italic>\n \n +\/+<\/jats:italic>\n <\/jats:sup>\n vs.\n Ghsr<\/jats:italic>\n \n \u2212\/\u2212<\/jats:italic>\n <\/jats:sup>\n : 91.75\u00a0\u00b1\u00a01.05% vs. 86.18\u00a0\u00b1\u00a01.13% of baseline BW,\n P<\/jats:italic>\n \u00a0<\u00a00.001)], gastrocnemius (T\u00a0+\u00a0G\u2010treated\n Ghsr<\/jats:italic>\n \n +\/+<\/jats:italic>\n <\/jats:sup>\n vs.\n Ghsr<\/jats:italic>\n \n \u2212\/\u2212<\/jats:italic>\n <\/jats:sup>\n : 96.9\u00a0\u00b1\u00a02.08% vs. 88.15\u00a0\u00b1\u00a01.78% of\n Ghsr<\/jats:italic>\n \n +\/+<\/jats:italic>\n <\/jats:sup>\n HK\u00a0+\u00a0V,\n P<\/jats:italic>\n \u00a0<\u00a00.001) and quadriceps muscle mass (T\u00a0+\u00a0G\u2010treated\n Ghsr<\/jats:italic>\n \n +\/+<\/jats:italic>\n <\/jats:sup>\n vs.\n Ghsr<\/jats:italic>\n \n \u2212\/\u2212<\/jats:italic>\n <\/jats:sup>\n : 96.12\u00a0\u00b1\u00a02.31% vs. 88.36\u00a0\u00b1\u00a01.94% of\n Ghsr<\/jats:italic>\n \n +\/+<\/jats:italic>\n <\/jats:sup>\n HK\u00a0+\u00a0V,\n P<\/jats:italic>\n \u00a0=\u00a00.01), and gastrocnemius type IIA (T\u00a0+\u00a0G\u2010treated\n Ghsr<\/jats:italic>\n \n +\/+<\/jats:italic>\n <\/jats:sup>\n vs.\n Ghsr<\/jats:italic>\n \n \u2212\/\u2212<\/jats:italic>\n <\/jats:sup>\n : 1250.49\u00a0\u00b1\u00a031.72 vs. 1017.62\u00a0\u00b1\u00a070.99\u00a0\u03bcm\n 2<\/jats:sup>\n ,\n P<\/jats:italic>\n \u00a0=\u00a00.027) and IIB fibre cross\u2010sectional area (T\u00a0+\u00a0G\u2010treated\n Ghsr<\/jats:italic>\n \n +\/+<\/jats:italic>\n <\/jats:sup>\n vs.\n Ghsr<\/jats:italic>\n \n \u2212\/\u2212<\/jats:italic>\n <\/jats:sup>\n : 2496.48\u00a0\u00b1\u00a0116.88 vs. 2183.04\u00a0\u00b1\u00a0103.43\u00a0\u03bcm\n 2<\/jats:sup>\n ,\n P<\/jats:italic>\n \u00a0=\u00a00.024).\n <\/jats:p>\n <\/jats:sec>\n \n Conclusions<\/jats:title>\n Growth hormone secretagogue receptor\u20101a mediates ghrelin's effects on attenuating LLC\u2010induced weakness but not muscle mass loss by modulating the autophagy\u2010lysosome pathway, mitophagy, and mitochondrial respiration.<\/jats:p>\n <\/jats:sec>","DOI":"10.1002\/jcsm.12743","type":"journal-article","created":{"date-parts":[[2021,7,15]],"date-time":"2021-07-15T06:56:54Z","timestamp":1626332214000},"page":"1280-1295","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":13,"title":["Growth hormone secretagogue receptor\u20101a mediates ghrelin's effects on attenuating tumour\u2010induced loss of muscle strength but not muscle mass"],"prefix":"10.1002","volume":"12","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-3142-3690","authenticated-orcid":false,"given":"Haiming","family":"Liu","sequence":"first","affiliation":[{"name":"Geriatric 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