{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,21]],"date-time":"2026-07-21T06:21:40Z","timestamp":1784614900180,"version":"3.55.0"},"reference-count":77,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2021,11,2]],"date-time":"2021-11-02T00:00:00Z","timestamp":1635811200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2021,11,2]],"date-time":"2021-11-02T00:00:00Z","timestamp":1635811200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Commun Biol"],"abstract":"<jats:title>Abstract<\/jats:title><jats:p>The gastric epithelium is often exposed to injurious elements and failure of appropriate healing predisposes to ulcers, hemorrhage, and\u00a0ultimately cancer. We examined the gastric function of CD36, a protein linked to disease and homeostasis. We used the tamoxifen model of gastric injury in mice null for <jats:italic>Cd36 (Cd36<\/jats:italic><jats:sup><jats:italic>\u2212\/\u2212<\/jats:italic><\/jats:sup>), with <jats:italic>Cd36<\/jats:italic> deletion in parietal cells (PC-<jats:italic>Cd36<\/jats:italic><jats:sup><jats:italic>\u2212\/\u2212<\/jats:italic><\/jats:sup>) or in endothelial cells (EC-<jats:italic>Cd36<\/jats:italic><jats:sup><jats:italic>\u2212\/\u2212<\/jats:italic><\/jats:sup>). CD36 expresses on corpus ECs, on PC basolateral membranes, and in gastrin and ghrelin cells. Stomachs of <jats:italic>Cd36<\/jats:italic><jats:sup><jats:italic>\u2212\/\u2212<\/jats:italic><\/jats:sup> mice have altered gland organization and secretion, more fibronectin, and inflammation. Tissue respiration and mitochondrial efficiency are reduced. Phospholipids increased and triglycerides decreased. Mucosal repair after injury is impaired in <jats:italic>Cd36<\/jats:italic><jats:sup><jats:italic>\u2212\/\u2212<\/jats:italic><\/jats:sup> and EC-<jats:italic>Cd36<\/jats:italic><jats:sup><jats:italic>\u2212\/\u2212<\/jats:italic><\/jats:sup>, not in PC-<jats:italic>Cd36<\/jats:italic><jats:sup><jats:italic>\u2212\/\u2212<\/jats:italic><\/jats:sup> mice, and is due to defect of progenitor differentiation to PCs, not of progenitor proliferation or mature PC dysfunction. Relevance to humans\u00a0is explored in the Vanderbilt BioVu using PrediXcan that links genetically-determined gene expression to clinical phenotypes, which\u00a0associates low <jats:italic>CD36<\/jats:italic> mRNA with gastritis, gastric ulcer, and gastro-intestinal hemorrhage. A CD36 variant predicted to disrupt an enhancer site associates (p\u2009&lt;\u200910<jats:sup>\u221217<\/jats:sup>) to death from gastro-intestinal hemorrhage in the UK Biobank. The findings support role of CD36 in gastric tissue repair, and\u00a0its deletion associated with chronic diseases that can predispose to malignancy.<\/jats:p>","DOI":"10.1038\/s42003-021-02765-z","type":"journal-article","created":{"date-parts":[[2021,11,2]],"date-time":"2021-11-02T11:02:39Z","timestamp":1635850959000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":20,"title":["CD36 maintains the gastric mucosa and associates with gastric disease"],"prefix":"10.1038","volume":"4","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-3949-3618","authenticated-orcid":false,"given":"Miriam","family":"Jacome-Sosa","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhi-Feng","family":"Miao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Vivek S.","family":"Peche","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Edward F.","family":"Morris","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ramkumar","family":"Narendran","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kathryn M.","family":"Pietka","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dmitri","family":"Samovski","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7253-6873","authenticated-orcid":false,"given":"Hei-Yong G.","family":"Lo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Terri","family":"Pietka","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andrea","family":"Varro","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Latisha","family":"Love-Gregory","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7833-2940","authenticated-orcid":false,"given":"James R.","family":"Goldenring","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7034-4536","authenticated-orcid":false,"given":"Ondrej","family":"Kuda","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4204-8734","authenticated-orcid":false,"given":"Eric R.","family":"Gamazon","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jason C.","family":"Mills","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6475-0877","authenticated-orcid":false,"given":"Nada A.","family":"Abumrad","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2021,11,2]]},"reference":[{"key":"2765_CR1","doi-asserted-by":"publisher","first-page":"452","DOI":"10.1097\/MOG.0000000000000308","volume":"32","author":"ML Schubert","year":"2016","unstructured":"Schubert, M. L. Gastric acid secretion. Curr. Opin. Gastroenterol. 32, 452\u2013460I (2016).","journal-title":"Curr. Opin. Gastroenterol."},{"key":"2765_CR2","doi-asserted-by":"publisher","first-page":"140","DOI":"10.1111\/obr.12474","volume":"18","author":"R Churm","year":"2017","unstructured":"Churm, R., Davies, J. S., Stephens, J. W. & Prior, S. L. Ghrelin function in human obesity and type 2 diabetes: a concise review. Obes. Rev. 18, 140\u2013148I (2017).","journal-title":"Obes. Rev."},{"key":"2765_CR3","doi-asserted-by":"crossref","unstructured":"Lv, Y., Liang, T., Wang, G. & Li, Z. Ghrelin, a gastrointestinal hormone, regulates energy balance and lipid metabolism. Biosci. Rep. 38, BSR20181061 (2018).","DOI":"10.1042\/BSR20181061"},{"key":"2765_CR4","doi-asserted-by":"publisher","first-page":"52","DOI":"10.2174\/1381612822666151119144458","volume":"22","author":"A Virdis","year":"2016","unstructured":"Virdis, A. et al. Human Ghrelin: A Gastric Hormone with Cardiovascular Properties. Curr. Pharm. Des. 22, 52\u201358I (2016).","journal-title":"Curr. Pharm. Des."},{"key":"2765_CR5","doi-asserted-by":"publisher","first-page":"1","DOI":"10.5115\/acb.2012.45.1.1","volume":"45","author":"P Cammisotto","year":"2012","unstructured":"Cammisotto, P. & Bendayan, M. A review on gastric leptin: the exocrine secretion of a gastric hormone. Anat. Cell Biol. 45, 1\u201316I (2012).","journal-title":"Anat. Cell Biol."},{"key":"2765_CR6","doi-asserted-by":"publisher","first-page":"259","DOI":"10.1053\/gast.2002.31385","volume":"122","author":"I Sobhani","year":"2002","unstructured":"Sobhani, I. et al. Vagal stimulation rapidly increases leptin secretion in human stomach. Gastroenterology 122, 259\u2013263I (2002).","journal-title":"Gastroenterology"},{"key":"2765_CR7","doi-asserted-by":"publisher","first-page":"79","DOI":"10.1053\/gast.2001.25490","volume":"121","author":"JC Hardwick","year":"2001","unstructured":"Hardwick, J. C., Van Den Brink, G. R., Offerhaus, G. J., Van Deventer, S. J. & Peppelenbosch, M. P. Leptin is a growth factor for colonic epithelial cells. Gastroenterology 121, 79\u201390I (2001).","journal-title":"Gastroenterology"},{"key":"2765_CR8","doi-asserted-by":"publisher","first-page":"928","DOI":"10.1007\/s00125-016-3898-3","volume":"59","author":"TH Meek","year":"2016","unstructured":"Meek, T. H. & Morton, G. J. The role of leptin in diabetes: metabolic effects. Diabetologia 59, 928\u2013932I (2016).","journal-title":"Diabetologia"},{"key":"2765_CR9","doi-asserted-by":"publisher","first-page":"269","DOI":"10.1016\/j.nut.2010.07.004","volume":"27","author":"SS Yarandi","year":"2011","unstructured":"Yarandi, S. S., Hebbar, G., Sauer, C. G., Cole, C. R. & Ziegler, T. R. Diverse roles of leptin in the gastrointestinal tract: modulation of motility, absorption, growth, and inflammation. Nutrition 27, 269\u2013275I (2011).","journal-title":"Nutrition"},{"key":"2765_CR10","doi-asserted-by":"publisher","first-page":"197","DOI":"10.1002\/cphy.c180035","volume":"10","author":"ML Schubert","year":"2019","unstructured":"Schubert, M. L. & Rehfeld, J. F. Gastric peptides-gastrin and somatostatin. Compr. Physiol. 10, 197\u2013228I (2019).","journal-title":"Compr. Physiol."},{"key":"2765_CR11","doi-asserted-by":"publisher","first-page":"17040I","DOI":"10.1038\/nrdp.2017.40","volume":"3","author":"R Green","year":"2017","unstructured":"Green, R. et al. Vitamin B(12) deficiency. Nat. Rev. Dis. Prim. 3, 17040I (2017).","journal-title":"Nat. Rev. Dis. Prim."},{"issue":"Suppl 1","key":"2765_CR12","doi-asserted-by":"publisher","first-page":"S37","DOI":"10.1097\/00004836-199112001-00006","volume":"13","author":"ND Yeomans","year":"1991","unstructured":"Yeomans, N. D. & Skeljo, M. V. Repair and healing of established gastric mucosal injury. J. Clin. Gastroenterol. 13(Suppl 1), S37\u201341I (1991).","journal-title":"J. Clin. Gastroenterol."},{"key":"2765_CR13","doi-asserted-by":"publisher","first-page":"625","DOI":"10.1016\/j.jcmgh.2016.05.005","volume":"2","author":"E Aihara","year":"2016","unstructured":"Aihara, E. et al. Epithelial Regeneration After Gastric Ulceration Causes Prolonged Cell-Type Alterations. Cell. Mol. Gastroenterol. Hepatol. 2, 625\u2013647I (2016).","journal-title":"Cell. Mol. Gastroenterol. Hepatol."},{"key":"2765_CR14","doi-asserted-by":"publisher","first-page":"3861","DOI":"10.1113\/JP275512","volume":"596","author":"AR Meyer","year":"2018","unstructured":"Meyer, A. R. & Goldenring, J. R. Injury, repair, inflammation and metaplasia in the stomach. J. Physiol. 596, 3861\u20133867I (2018).","journal-title":"J. Physiol."},{"key":"2765_CR15","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1038\/nature21028","volume":"542","author":"BW Wong","year":"2017","unstructured":"Wong, B. W. et al. The role of fatty acid beta-oxidation in lymphangiogenesis. Nature 542, 49\u201354I (2017).","journal-title":"Nature"},{"key":"2765_CR16","doi-asserted-by":"publisher","first-page":"689","DOI":"10.1016\/j.molcel.2018.01.010","volume":"69","author":"J Xiong","year":"2018","unstructured":"Xiong, J. et al. A metabolic basis for endothelial-to-mesenchymal transition. Mol. Cell 69, 689\u2013698 (2018). e687I.","journal-title":"Mol. Cell"},{"key":"2765_CR17","doi-asserted-by":"publisher","first-page":"302","DOI":"10.1038\/nature14190","volume":"517","author":"A Efeyan","year":"2015","unstructured":"Efeyan, A., Comb, W. C. & Sabatini, D. M. Nutrient-sensing mechanisms and pathways. Nature 517, 302\u2013310I (2015).","journal-title":"Nature"},{"key":"2765_CR18","doi-asserted-by":"publisher","first-page":"769","DOI":"10.1016\/j.stem.2018.04.001","volume":"22","author":"MM Mihaylova","year":"2018","unstructured":"Mihaylova, M. M. et al. Fasting activates fatty acid oxidation to enhance intestinal stem cell function during homeostasis and aging. cell stem cell 22, 769\u2013778 (2018). e764I.","journal-title":"cell stem cell"},{"key":"2765_CR19","doi-asserted-by":"publisher","first-page":"985","DOI":"10.1053\/j.gastro.2019.11.031","volume":"158","author":"L Chen","year":"2020","unstructured":"Chen, L. et al. HNF4 Regulates Fatty Acid Oxidation and Is Required for Renewal of Intestinal Stem Cells in Mice. Gastroenterology 158, 985\u2013999 (2020). e989I.","journal-title":"Gastroenterology"},{"key":"2765_CR20","doi-asserted-by":"publisher","first-page":"E916","DOI":"10.1152\/ajpendo.2001.281.5.E916","volume":"281","author":"M Chen","year":"2001","unstructured":"Chen, M., Yang, Y., Braunstein, E., Georgeson, K. E. & Harmon, C. M. Gut expression and regulation of FAT\/CD36: possible role in fatty acid transport in rat enterocytes. Am. J. Physiol. Endocrinol. Metab. 281, E916\u2013923I (2001).","journal-title":"Am. J. Physiol. Endocrinol. Metab."},{"key":"2765_CR21","doi-asserted-by":"publisher","first-page":"13687","DOI":"10.1073\/pnas.231332398","volume":"98","author":"JC Mills","year":"2001","unstructured":"Mills, J. C. et al. A molecular profile of the mouse gastric parietal cell with and without exposure to Helicobacter pylori. Proc. Natl Acad. Sci. USA 98, 13687\u201313692I (2001).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2765_CR22","doi-asserted-by":"publisher","first-page":"353","DOI":"10.2337\/db14-0582","volume":"64","author":"D Samovski","year":"2015","unstructured":"Samovski, D. et al. Regulation of AMPK activation by CD36 links fatty acid uptake to beta-oxidation. Diabetes 64, 353\u2013359I (2015).","journal-title":"Diabetes"},{"key":"2765_CR23","doi-asserted-by":"publisher","first-page":"621","DOI":"10.1038\/nri3515","volume":"13","author":"J Canton","year":"2013","unstructured":"Canton, J., Neculai, D. & Grinstein, S. Scavenger receptors in homeostasis and immunity. Nat. Rev. Immunol. 13, 621\u2013634I (2013).","journal-title":"Nat. Rev. Immunol."},{"key":"2765_CR24","doi-asserted-by":"publisher","first-page":"82","DOI":"10.1016\/j.jcmgh.2016.09.001","volume":"3","author":"V Cifarelli","year":"2017","unstructured":"Cifarelli, V. et al. CD36 deficiency impairs the small intestinal barrier and induces subclinical inflammation in mice. Cell. Mol. Gastroenterol. Hepatol. 3, 82\u201398I (2017).","journal-title":"Cell. Mol. Gastroenterol. Hepatol."},{"key":"2765_CR25","doi-asserted-by":"publisher","first-page":"1061","DOI":"10.1152\/physrev.00019.2011","volume":"92","author":"NA Abumrad","year":"2012","unstructured":"Abumrad, N. A. & Davidson, N. O. Role of the gut in lipid homeostasis. Physiol. Rev. 92, 1061\u20131085I (2012).","journal-title":"Physiol. Rev."},{"key":"2765_CR26","doi-asserted-by":"publisher","first-page":"1191","DOI":"10.1096\/fj.12-217703","volume":"27","author":"S Sundaresan","year":"2013","unstructured":"Sundaresan, S. et al. CD36-dependent signaling mediates fatty acid-induced gut release of secretin and cholecystokinin. Faseb j. 27, 1191\u20131202I (2013).","journal-title":"Faseb j."},{"key":"2765_CR27","doi-asserted-by":"publisher","first-page":"25201","DOI":"10.1074\/jbc.M111.233551","volume":"286","author":"TT Tran","year":"2011","unstructured":"Tran, T. T. et al. Luminal lipid regulates CD36 levels and downstream signaling to stimulate chylomicron synthesis. J. Biol. Chem. 286, 25201\u201325210I (2011).","journal-title":"J. Biol. Chem."},{"key":"2765_CR28","doi-asserted-by":"publisher","first-page":"154","DOI":"10.1101\/gad.285684.116","volume":"31","author":"HG Lo","year":"2017","unstructured":"Lo, H. G. et al. A single transcription factor is sufficient to induce and maintain secretory cell architecture. Genes Dev. 31, 154\u2013171I (2017).","journal-title":"Genes Dev."},{"key":"2765_CR29","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1152\/physiolgenomics.90408.2008","volume":"37","author":"BJ Capoccia","year":"2009","unstructured":"Capoccia, B. J., Huh, W. J. & Mills, J. C. How form follows functional genomics: gene expression profiling gastric epithelial cells with a particular discourse on the parietal cell. Physiol. Genomics 37, 67\u201378I (2009).","journal-title":"Physiol. Genomics"},{"key":"2765_CR30","doi-asserted-by":"publisher","first-page":"368","DOI":"10.1111\/j.1432-1033.1996.0368z.x","volume":"238","author":"H Poirier","year":"1996","unstructured":"Poirier, H., Degrace, P., Niot, I., Bernard, A. & Besnard, P. Localization and regulation of the putative membrane fatty-acid transporter (FAT) in the small intestine. Comparison with fatty acid-binding proteins (FABP). Eur. J. Biochem 238, 368\u2013373I (1996).","journal-title":"Eur. J. Biochem"},{"key":"2765_CR31","first-page":"199","volume":"22","author":"PG Cammisotto","year":"2007","unstructured":"Cammisotto, P. G. & Bendayan, M. Leptin secretion by white adipose tissue and gastric mucosa. Histol. Histopathol. 22, 199\u2013210I (2007).","journal-title":"Histol. Histopathol."},{"key":"2765_CR32","doi-asserted-by":"publisher","first-page":"790","DOI":"10.1038\/29547","volume":"394","author":"A Bado","year":"1998","unstructured":"Bado, A. et al. The stomach is a source of leptin. Nature 394, 790\u2013793I (1998).","journal-title":"Nature"},{"key":"2765_CR33","doi-asserted-by":"publisher","first-page":"789","DOI":"10.1038\/sj.ijo.0801228","volume":"24","author":"S Cinti","year":"2000","unstructured":"Cinti, S. et al. Secretory granules of endocrine and chief cells of human stomach mucosa contain leptin. Int J. Obes. Relat. Metab. Disord. 24, 789\u2013793I (2000).","journal-title":"Int J. Obes. Relat. Metab. Disord."},{"key":"2765_CR34","doi-asserted-by":"publisher","first-page":"178","DOI":"10.1136\/gut.47.2.178","volume":"47","author":"I Sobhani","year":"2000","unstructured":"Sobhani, I. et al. Leptin secretion and leptin receptor in the human stomach. Gut 47, 178\u2013183I (2000).","journal-title":"Gut"},{"key":"2765_CR35","doi-asserted-by":"publisher","first-page":"159","DOI":"10.1007\/s11154-016-9355-2","volume":"17","author":"P Besnard","year":"2016","unstructured":"Besnard, P. Lipids and obesity: Also a matter of taste? Rev. Endocr. Metab. Disord. 17, 159\u2013170I (2016).","journal-title":"Rev. Endocr. Metab. Disord."},{"key":"2765_CR36","doi-asserted-by":"publisher","first-page":"1339","DOI":"10.1002\/cphy.c130055","volume":"4","author":"KN Browning","year":"2014","unstructured":"Browning, K. N. & Travagli, R. A. Central nervous system control of gastrointestinal motility and secretion and modulation of gastrointestinal functions. Compr. Physiol. 4, 1339\u20131368I (2014).","journal-title":"Compr. Physiol."},{"key":"2765_CR37","doi-asserted-by":"publisher","first-page":"3177","DOI":"10.1172\/JCI25299","volume":"115","author":"F Laugerette","year":"2005","unstructured":"Laugerette, F. et al. CD36 involvement in orosensory detection of dietary lipids, spontaneous fat preference, and digestive secretions. J. Clin. Investig. 115, 3177\u20133184I (2005).","journal-title":"J. Clin. Investig."},{"key":"2765_CR38","doi-asserted-by":"publisher","first-page":"573","DOI":"10.1152\/physrev.00016.2019","volume":"100","author":"AC Engevik","year":"2020","unstructured":"Engevik, A. C., Kaji, I. & Goldenring, J. R. The Physiology of the Gastric Parietal Cell. Physiol. Rev. 100, 573\u2013602I (2020).","journal-title":"Physiol. Rev."},{"key":"2765_CR39","doi-asserted-by":"publisher","first-page":"G521","DOI":"10.1152\/ajpgi.00105.2004","volume":"289","author":"G Xie","year":"2005","unstructured":"Xie, G., Drachenberg, C., Yamada, M., Wess, J. & Raufman, J. P. Cholinergic agonist-induced pepsinogen secretion from murine gastric chief cells is mediated by M1 and M3 muscarinic receptors. Am. J. Physiol. Gastrointest. Liver Physiol. 289, G521\u2013529I (2005).","journal-title":"Am. J. Physiol. Gastrointest. Liver Physiol."},{"key":"2765_CR40","doi-asserted-by":"publisher","first-page":"278I","DOI":"10.3389\/fphar.2016.00278","volume":"7","author":"K Takeuchi","year":"2016","unstructured":"Takeuchi, K., Endoh, T., Hayashi, S. & Aihara, T. Activation of Muscarinic Acetylcholine Receptor Subtype 4 Is Essential for Cholinergic Stimulation of Gastric Acid Secretion: Relation to D Cell\/Somatostatin. Front Pharm. 7, 278I (2016).","journal-title":"Front Pharm."},{"key":"2765_CR41","doi-asserted-by":"publisher","first-page":"21","DOI":"10.1053\/j.gastro.2011.09.050","volume":"142","author":"WJ Huh","year":"2012","unstructured":"Huh, W. J. et al. Tamoxifen induces rapid, reversible atrophy, and metaplasia in mouse stomach. Gastroenterology 142, 21\u201324 (2012). e27I.","journal-title":"Gastroenterology"},{"key":"2765_CR42","doi-asserted-by":"publisher","first-page":"329","DOI":"10.1007\/978-1-4939-3603-8_28","volume":"1422","author":"JB Saenz","year":"2016","unstructured":"Saenz, J. B., Burclaff, J. & Mills, J. C. Modeling Murine Gastric Metaplasia Through Tamoxifen-Induced Acute Parietal Cell Loss. Methods Mol. Biol. 1422, 329\u2013339I (2016).","journal-title":"Methods Mol. Biol."},{"key":"2765_CR43","doi-asserted-by":"publisher","first-page":"582","DOI":"10.1053\/j.gastro.2004.05.029","volume":"127","author":"S Nomura","year":"2004","unstructured":"Nomura, S. et al. Spasmolytic polypeptide expressing metaplasia to preneoplasia in H. felis-infected mice. Gastroenterology 127, 582\u2013594I (2004).","journal-title":"Gastroenterology"},{"key":"2765_CR44","doi-asserted-by":"publisher","first-page":"53","DOI":"10.1038\/nature17173","volume":"531","author":"S Beyaz","year":"2016","unstructured":"Beyaz, S. et al. High-fat diet enhances stemness and tumorigenicity of intestinal progenitors. Nature 531, 53\u201358I (2016).","journal-title":"Nature"},{"key":"2765_CR45","doi-asserted-by":"publisher","first-page":"2144","DOI":"10.1016\/j.celrep.2017.08.029","volume":"20","author":"M Knobloch","year":"2017","unstructured":"Knobloch, M. et al. A Fatty Acid Oxidation-Dependent Metabolic Shift Regulates Adult Neural Stem Cell Activity. Cell Rep. 20, 2144\u20132155I (2017).","journal-title":"Cell Rep."},{"key":"2765_CR46","doi-asserted-by":"publisher","first-page":"4329","DOI":"10.1172\/JCI99315","volume":"128","author":"NH Son","year":"2018","unstructured":"Son, N. H. et al. Endothelial cell CD36 optimizes tissue fatty acid uptake. The. J. Clin. Investig. 128, 4329\u20134342I (2018).","journal-title":"J. Clin. Investig."},{"key":"2765_CR47","doi-asserted-by":"crossref","unstructured":"Miller, M. J., Cusmano-Ozog, K., Oglesbee, D. & Young, S. Laboratory analysis of acylcarnitines, 2020 update: a technical standard of the American College of Medical Genetics and Genomics (ACMG). Genet Med. 23, 249\u2013258 (2020).","DOI":"10.1038\/s41436-020-00990-1"},{"key":"2765_CR48","doi-asserted-by":"publisher","first-page":"45","DOI":"10.1016\/j.cmet.2007.10.013","volume":"7","author":"TR Koves","year":"2008","unstructured":"Koves, T. R. et al. Mitochondrial overload and incomplete fatty acid oxidation contribute to skeletal muscle insulin resistance. Cell Metab. 7, 45\u201356I (2008).","journal-title":"Cell Metab."},{"key":"2765_CR49","doi-asserted-by":"publisher","first-page":"2I","DOI":"10.1186\/1476-511X-5-2","volume":"5","author":"HJ Lee","year":"2006","unstructured":"Lee, H. J., Mayette, J., Rapoport, S. I. & Bazinet, R. P. Selective remodeling of cardiolipin fatty acids in the aged rat heart. Lipids Health Dis. 5, 2I (2006).","journal-title":"Lipids Health Dis."},{"key":"2765_CR50","doi-asserted-by":"publisher","first-page":"6417","DOI":"10.1021\/bi7004015","volume":"46","author":"X Han","year":"2007","unstructured":"Han, X. et al. Alterations in myocardial cardiolipin content and composition occur at the very earliest stages of diabetes: a shotgun lipidomics study. Biochemistry 46, 6417\u20136428I (2007).","journal-title":"Biochemistry"},{"key":"2765_CR51","doi-asserted-by":"publisher","first-page":"456","DOI":"10.1136\/gut.34.4.456","volume":"34","author":"G Nardone","year":"1993","unstructured":"Nardone, G., Laccetti, P., Civiletti, C. & Budillon, G. Phospholipid composition of human gastric mucosa: a study of endoscopic biopsy specimens. Gut 34, 456\u2013460I (1993).","journal-title":"Gut"},{"key":"2765_CR52","doi-asserted-by":"publisher","first-page":"956","DOI":"10.1038\/s41588-018-0154-4","volume":"50","author":"ER Gamazon","year":"2018","unstructured":"Gamazon, E. R. et al. Using an atlas of gene regulation across 44 human tissues to inform complex disease- and trait-associated variation. Nat. Genet 50, 956\u2013967I (2018).","journal-title":"Nat. Genet"},{"key":"2765_CR53","doi-asserted-by":"publisher","first-page":"204","DOI":"10.1038\/nature24277","volume":"550","author":"A Battle","year":"2017","unstructured":"Battle, A., Brown, C. D., Engelhardt, B. E. & Montgomery, S. B. Genetic effects on gene expression across human tissues. Nature 550, 204\u2013213I (2017).","journal-title":"Nature"},{"key":"2765_CR54","doi-asserted-by":"publisher","first-page":"98","DOI":"10.1038\/s41591-019-0705-y","volume":"26","author":"G Unlu","year":"2020","unstructured":"Unlu, G. et al. Phenome-based approach identifies RIC1-linked Mendelian syndrome through zebrafish models, biobank associations and clinical studies. Nat. Med 26, 98\u2013109I (2020).","journal-title":"Nat. Med"},{"key":"2765_CR55","doi-asserted-by":"publisher","first-page":"203","DOI":"10.1038\/s41586-018-0579-z","volume":"562","author":"C Bycroft","year":"2018","unstructured":"Bycroft, C. et al. The UK Biobank resource with deep phenotyping and genomic data. Nature 562, 203\u2013209I (2018).","journal-title":"Nature"},{"key":"2765_CR56","doi-asserted-by":"publisher","first-page":"317","DOI":"10.1038\/nature14248","volume":"518","author":"A Kundaje","year":"2015","unstructured":"Kundaje, A. et al. Integrative analysis of 111 reference human epigenomes. Nature 518, 317\u2013330I (2015).","journal-title":"Nature"},{"key":"2765_CR57","doi-asserted-by":"publisher","first-page":"2176","DOI":"10.1194\/jlr.P065250","volume":"57","author":"L Love-Gregory","year":"2016","unstructured":"Love-Gregory, L. et al. Higher chylomicron remnants and LDL particle numbers associate with CD36 SNPs and DNA methylation sites that reduce CD36. J. Lipid Res 57, 2176\u20132184I (2016).","journal-title":"J. Lipid Res"},{"key":"2765_CR58","doi-asserted-by":"publisher","first-page":"1995s","DOI":"10.1093\/jn\/nxy201","volume":"148","author":"LH Allen","year":"2018","unstructured":"Allen, L. H. et al. Biomarkers of Nutrition for Development (BOND): Vitamin B-12 Review. J. Nutr. 148, 1995s\u20132027sI (2018).","journal-title":"J. Nutr."},{"key":"2765_CR59","doi-asserted-by":"publisher","first-page":"340","DOI":"10.1159\/000485734","volume":"97","author":"K Haruma","year":"2018","unstructured":"Haruma, K. et al. Old and New Gut Hormone, Gastrin and Acid Suppressive Therapy. Digestion 97, 340\u2013344I (2018).","journal-title":"Digestion"},{"key":"2765_CR60","doi-asserted-by":"publisher","first-page":"2951","DOI":"10.1113\/jphysiol.2014.272435","volume":"592","author":"R Dimaline","year":"2014","unstructured":"Dimaline, R. & Varro, A. Novel roles of gastrin. J. Physiol. 592, 2951\u20132958I (2014).","journal-title":"J. Physiol."},{"key":"2765_CR61","doi-asserted-by":"publisher","first-page":"37I","DOI":"10.1186\/1471-230X-5-37","volume":"5","author":"A Hacioglu","year":"2005","unstructured":"Hacioglu, A., Algin, C., Pasaoglu, O., Pasaoglu, E. & Kanbak, G. Protective effect of leptin against ischemia-reperfusion injury in the rat small intestine. BMC Gastroenterol. 5, 37I (2005).","journal-title":"BMC Gastroenterol."},{"key":"2765_CR62","doi-asserted-by":"publisher","first-page":"4154","DOI":"10.3748\/wjg.v11.i27.4154","volume":"11","author":"EO Adeyemi","year":"2005","unstructured":"Adeyemi, E. O. et al. Mechanisms of action of leptin in preventing gastric ulcer. World J. Gastroenterol. 11, 4154\u20134160I (2005).","journal-title":"World J. Gastroenterol."},{"key":"2765_CR63","doi-asserted-by":"publisher","first-page":"23502","DOI":"10.1074\/jbc.M111.315358","volume":"287","author":"JT McFarlan","year":"2012","unstructured":"McFarlan, J. T. et al. In vivo, fatty acid translocase (CD36) critically regulates skeletal muscle fuel selection, exercise performance, and training-induced adaptation of fatty acid oxidation. J. Biol. Chem. 287, 23502\u201323516I (2012).","journal-title":"J. Biol. Chem."},{"key":"2765_CR64","doi-asserted-by":"publisher","first-page":"4415","DOI":"10.1113\/jphysiol.2012.238451","volume":"591","author":"Y Yoshida","year":"2013","unstructured":"Yoshida, Y. et al. Exercise- and training-induced upregulation of skeletal muscle fatty acid oxidation are not solely dependent on mitochondrial machinery and biogenesis. J. Physiol. 591, 4415\u20134426I (2013).","journal-title":"J. Physiol."},{"key":"2765_CR65","doi-asserted-by":"publisher","first-page":"14317","DOI":"10.1074\/jbc.M706478200","volume":"283","author":"Z Nahl\u00e9","year":"2008","unstructured":"Nahl\u00e9, Z. et al. CD36-dependent regulation of muscle FoxO1 and PDK4 in the PPAR delta\/beta-mediated adaptation to metabolic stress. J. Biol. Chem. 283, 14317\u201314326I (2008).","journal-title":"J. Biol. Chem."},{"key":"2765_CR66","doi-asserted-by":"crossref","unstructured":"Willet, S. G. et al. Regenerative proliferation of differentiated cells by mTORC1-dependent paligenosis. EMBO J 37, e98311 (2018).","DOI":"10.15252\/embj.201798311"},{"key":"2765_CR67","doi-asserted-by":"crossref","unstructured":"Harayama, T. & Shimizu, T. Roles of polyunsaturated fatty acids, from mediators to membranes. J. Lipid. Res. 61, 1150\u20131160I (2020).","DOI":"10.1194\/jlr.R120000800"},{"key":"2765_CR68","doi-asserted-by":"publisher","first-page":"4282","DOI":"10.1074\/jbc.AW118.003229","volume":"294","author":"DR Voelker","year":"2019","unstructured":"Voelker, D. R. & Numata, M. Phospholipid regulation of innate immunity and respiratory viral infection. J. Biol. Chem. 294, 4282\u20134289I (2019).","journal-title":"J. Biol. Chem."},{"key":"2765_CR69","doi-asserted-by":"publisher","first-page":"80","DOI":"10.1016\/j.stem.2008.11.011","volume":"4","author":"K Cui","year":"2009","unstructured":"Cui, K. et al. Chromatin signatures in multipotent human hematopoietic stem cells indicate the fate of bivalent genes during differentiation. cell stem cell 4, 80\u201393I (2009).","journal-title":"cell stem cell"},{"key":"2765_CR70","doi-asserted-by":"publisher","first-page":"90","DOI":"10.1038\/nrm.2016.140","volume":"18","author":"BR Sabari","year":"2017","unstructured":"Sabari, B. R., Zhang, D., Allis, C. D. & Zhao, Y. Metabolic regulation of gene expression through histone acylations. Nat. Rev. Mol. Cell Biol. 18, 90\u2013101I (2017).","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"2765_CR71","doi-asserted-by":"publisher","first-page":"1695","DOI":"10.1093\/hmg\/ddn060","volume":"17","author":"L Love-Gregory","year":"2008","unstructured":"Love-Gregory, L. et al. Variants in the CD36 gene associate with the metabolic syndrome and high-density lipoprotein cholesterol. Hum. Mol. Genet 17, 1695\u20131704I (2008).","journal-title":"Hum. Mol. Genet"},{"key":"2765_CR72","doi-asserted-by":"publisher","first-page":"2197","DOI":"10.1093\/hmg\/ddh233","volume":"13","author":"X Ma","year":"2004","unstructured":"Ma, X. et al. A common haplotype at the CD36 locus is associated with high free fatty acid levels and increased cardiovascular risk in Caucasians. Hum. Mol. Genet 13, 2197\u20132205I (2004).","journal-title":"Hum. Mol. Genet"},{"key":"2765_CR73","doi-asserted-by":"publisher","first-page":"1179","DOI":"10.1053\/j.gastro.2012.08.002","volume":"143","author":"AF Peery","year":"2012","unstructured":"Peery, A. F. et al. Burden of gastrointestinal disease in the United States: 2012 update. Gastroenterology 143, 1179\u20131187 (2012). e1173I.","journal-title":"Gastroenterology"},{"key":"2765_CR74","doi-asserted-by":"publisher","first-page":"300","DOI":"10.2337\/db19-0494","volume":"69","author":"V Paluchova","year":"2020","unstructured":"Paluchova, V. et al. Lipokine 5-PAHSA Is Regulated by Adipose Triglyceride Lipase and Primes Adipocytes for De Novo Lipogenesis in Mice. Diabetes 69, 300\u2013312I (2020).","journal-title":"Diabetes"},{"key":"2765_CR75","doi-asserted-by":"crossref","unstructured":"Cajka, T., Smilowitz, J. T. & Fiehn, O. Validating Quantitative Untargeted Lipidomics Across Nine Liquid Chromatography-High-Resolution Mass Spectrometry Platforms. Anal. Chem. 89, 12360-\u20132368I (2017).","DOI":"10.1021\/acs.analchem.7b03404"},{"key":"2765_CR76","doi-asserted-by":"publisher","first-page":"211","DOI":"10.1242\/dev.02700","volume":"134","author":"VG Ramsey","year":"2007","unstructured":"Ramsey, V. G. et al. The maturation of mucus-secreting gastric epithelial progenitors into digestive-enzyme secreting zymogenic cells requires Mist1. Development 134, 211\u2013222I (2007).","journal-title":"Development"},{"key":"2765_CR77","doi-asserted-by":"publisher","first-page":"1091","DOI":"10.1038\/ng.3367","volume":"47","author":"ER Gamazon","year":"2015","unstructured":"Gamazon, E. R. et al. A gene-based association method for mapping traits using reference transcriptome data. Nat. Genet 47, 1091\u20131098I (2015).","journal-title":"Nat. Genet"}],"container-title":["Communications Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s42003-021-02765-z.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s42003-021-02765-z","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s42003-021-02765-z.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,12,3]],"date-time":"2022-12-03T19:50:44Z","timestamp":1670097044000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s42003-021-02765-z"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,11,2]]},"references-count":77,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2021,12]]}},"alternative-id":["2765"],"URL":"https:\/\/doi.org\/10.1038\/s42003-021-02765-z","relation":{},"ISSN":["2399-3642"],"issn-type":[{"value":"2399-3642","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,11,2]]},"assertion":[{"value":"29 March 2021","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"6 October 2021","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"2 November 2021","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"1247"}}