{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,1,27]],"date-time":"2026-01-27T10:17:09Z","timestamp":1769509029017,"version":"3.49.0"},"reference-count":69,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2020,10,26]],"date-time":"2020-10-26T00:00:00Z","timestamp":1603670400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2020,10,26]],"date-time":"2020-10-26T00:00:00Z","timestamp":1603670400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100009207","name":"Agrifutures Australia","doi-asserted-by":"crossref","award":["PRJ-010413"],"award-info":[{"award-number":["PRJ-010413"]}],"id":[{"id":"10.13039\/501100009207","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title><jats:p>Inborn errors of metabolism are genetic conditions that can disrupt intermediary metabolic pathways and cause defective absorption and metabolism of dietary nutrients. In an Australian Kelpie breeding population, 17 puppies presented with intestinal lipid malabsorption. Juvenile dogs exhibited stunted postnatal growth, steatorrhea, abdominal distension and a wiry coat. Using genome-wide association analysis, an associated locus on CFA28 (P<jats:sub>raw<\/jats:sub>\u2009=\u20092.87E<jats:sup>\u221206<\/jats:sup>) was discovered and validated in a closely related population (P<jats:sub>raw<\/jats:sub>\u2009=\u20091.75E<jats:sup>\u221245<\/jats:sup>). A 103.3\u00a0kb deletion NC_006610.3CFA28:g.23380074_23483377del, containing genes Acyl-CoA Synthetase Long Chain Family Member 5 (<jats:italic>ACSL5<\/jats:italic>) and Zinc Finger DHHC-Type Containing 6 (<jats:italic>ZDHHC6<\/jats:italic>), was characterised using whole transcriptomic data. Whole transcriptomic sequencing revealed no expression of <jats:italic>ACSL5<\/jats:italic> and disrupted splicing of <jats:italic>ZDHHC6<\/jats:italic> in jejunal tissue of affected Kelpies. The <jats:italic>ACSL5<\/jats:italic> gene plays a key role in long chain fatty acid absorption, a phenotype similar to that of our affected Kelpies has been observed in a knockout mouse model. A PCR-based diagnostic test was developed and confirmed fully penetrant autosomal recessive mode of inheritance. We conclude the structural variant causing a deletion of the <jats:italic>ACSL5<\/jats:italic> gene is the most likely cause for intestinal lipid malabsorption in the Australian Kelpie.<\/jats:p>","DOI":"10.1038\/s41598-020-75243-x","type":"journal-article","created":{"date-parts":[[2020,10,26]],"date-time":"2020-10-26T11:03:40Z","timestamp":1603710220000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":6,"title":["A large deletion on CFA28 omitting ACSL5 gene is associated with intestinal lipid malabsorption in the Australian Kelpie dog breed"],"prefix":"10.1038","volume":"10","author":[{"given":"Mitchell J.","family":"O\u2019Brien","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Niek J.","family":"Beijerink","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Mandy","family":"Sansom","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Sarah W.","family":"Thornton","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Tracy","family":"Chew","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Claire M.","family":"Wade","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"297","published-online":{"date-parts":[[2020,10,26]]},"reference":[{"key":"75243_CR1","doi-asserted-by":"publisher","first-page":"1411","DOI":"10.1016\/j.bbadis.2012.02.010","volume":"1822","author":"PA Watkins","year":"2012","unstructured":"Watkins, P. A. & Ellis, J. M. Peroxisomal acyl-CoA synthetases. Biochim. Biophys. Acta BBA Mol. Basis Disease 1822, 1411\u20131420. https:\/\/doi.org\/10.1016\/j.bbadis.2012.02.010 (2012).","journal-title":"Biochim. Biophys. Acta BBA Mol. Basis Disease"},{"key":"75243_CR2","doi-asserted-by":"publisher","first-page":"2736","DOI":"10.1194\/jlr.M700378-JLR200","volume":"48","author":"PA Watkins","year":"2007","unstructured":"Watkins, P. A., Maiguel, D., Jia, Z. & Pevsner, J. Evidence for 26 distinct acyl-coenzyme A synthetase genes in the human genome. J. Lipid Res. 48, 2736\u20132750. https:\/\/doi.org\/10.1194\/jlr.M700378-JLR200 (2007).","journal-title":"J. Lipid Res."},{"key":"75243_CR3","doi-asserted-by":"publisher","first-page":"465","DOI":"10.2217\/17460875.2.4.465","volume":"2","author":"DG Mashek","year":"2007","unstructured":"Mashek, D. G., Li, L. O. & Coleman, R. A. Long-chain acyl-CoA synthetases and fatty acid channeling. Future Lipidol. 2, 465\u2013476. https:\/\/doi.org\/10.2217\/17460875.2.4.465 (2007).","journal-title":"Future Lipidol."},{"key":"75243_CR4","doi-asserted-by":"publisher","first-page":"6","DOI":"10.1186\/s40575-015-0017-6","volume":"2","author":"ER Arnott","year":"2015","unstructured":"Arnott, E. R. et al. Strong selection for behavioural resilience in Australian stock working dogs identified by selective sweep analysis. Canine Genet. Epidemiol. 2, 6. https:\/\/doi.org\/10.1186\/s40575-015-0017-6 (2015).","journal-title":"Canine Genet. Epidemiol."},{"key":"75243_CR5","doi-asserted-by":"publisher","DOI":"10.3390\/genes10050337","author":"T Chew","year":"2019","unstructured":"Chew, T., Willet, C. E., Haase, B. & Wade, C. M. Genomic characterization of external morphology traits in kelpies does not support common ancestry with the Australian Dingo. Genes. https:\/\/doi.org\/10.3390\/genes10050337 (2019).","journal-title":"Genes."},{"key":"75243_CR6","doi-asserted-by":"publisher","first-page":"418","DOI":"10.1186\/s12917-018-1752-1","volume":"14","author":"JC Fyfe","year":"2018","unstructured":"Fyfe, J. C. et al. Inherited selective cobalamin malabsorption in Komondor dogs associated with a CUBN splice site variant. BMC Vet. Res. 14, 418. https:\/\/doi.org\/10.1186\/s12917-018-1752-1 (2018).","journal-title":"BMC Vet. Res."},{"key":"75243_CR7","doi-asserted-by":"publisher","first-page":"390","DOI":"10.1016\/j.ymgme.2013.05.006","volume":"109","author":"JC Fyfe","year":"2013","unstructured":"Fyfe, J. C. et al. An exon 53 frameshift mutation in CUBN abrogates cubam function and causes Imerslund-Gr\u00e4sbeck syndrome in dogs. Mol. Genet. Metab. 109, 390\u2013396. https:\/\/doi.org\/10.1016\/j.ymgme.2013.05.006 (2013).","journal-title":"Mol. Genet. Metab."},{"key":"75243_CR8","doi-asserted-by":"publisher","first-page":"356","DOI":"10.1111\/jvim.12284","volume":"28","author":"JC Fyfe","year":"2014","unstructured":"Fyfe, J. C., Hemker, S. L., Venta, P. J., Stebbing, B. & Giger, U. Selective intestinal cobalamin malabsorption with proteinuria (Imerslund-Gr\u00e4sbeck syndrome) in juvenile Beagles. J. Vet. Intern. Med. 28, 356\u2013362. https:\/\/doi.org\/10.1111\/jvim.12284 (2014).","journal-title":"J. Vet. Intern. Med."},{"key":"75243_CR9","first-page":"1029","volume":"56","author":"AJ Gold","year":"2015","unstructured":"Gold, A. J., Scott, M. A. & Fyfe, J. C. Failure to thrive and life-threatening complications due to inherited selective cobalamin malabsorption effectively managed in a juvenile Australian shepherd dog. Can. Vet. J. 56, 1029\u20131034 (2015).","journal-title":"Can. Vet. J."},{"key":"75243_CR10","doi-asserted-by":"publisher","first-page":"1447","DOI":"10.1182\/blood-2005-03-1197","volume":"106","author":"Q He","year":"2005","unstructured":"He, Q. et al. Amnionless function is required for cubilin brush-border expression and intrinsic factor-cobalamin (vitamin B12) absorption in vivo. Blood 106, 1447\u20131453. https:\/\/doi.org\/10.1182\/blood-2005-03-1197 (2005).","journal-title":"Blood"},{"key":"75243_CR11","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1111\/jvim.15638","volume":"34","author":"S Kather","year":"2020","unstructured":"Kather, S., Gr\u00fctzner, N., Kook, P. H., Dengler, F. & Heilmann, R. M. Review of cobalamin status and disorders of cobalamin metabolism in dogs. J. Vet. Intern. Med. 34, 13\u201328. https:\/\/doi.org\/10.1111\/jvim.15638 (2020).","journal-title":"J. Vet. Intern. Med."},{"key":"75243_CR12","doi-asserted-by":"publisher","DOI":"10.1186\/s13742-015-0047-8","author":"CC Chang","year":"2015","unstructured":"Chang, C. C. et al. Second-generation PLINK: Rising to the challenge of larger and richer datasets. GigaScience. https:\/\/doi.org\/10.1186\/s13742-015-0047-8 (2015).","journal-title":"GigaScience."},{"key":"75243_CR13","doi-asserted-by":"publisher","first-page":"1937","DOI":"10.1093\/bioinformatics\/bty036","volume":"34","author":"T K\u00f5ressaar","year":"2018","unstructured":"K\u00f5ressaar, T. et al. Primer3_masker: Integrating masking of template sequence with primer design software. Bioinformatics 34, 1937\u20131938. https:\/\/doi.org\/10.1093\/bioinformatics\/bty036 (2018).","journal-title":"Bioinformatics"},{"key":"75243_CR14","doi-asserted-by":"publisher","first-page":"1289","DOI":"10.1093\/bioinformatics\/btm091","volume":"23","author":"T Koressaar","year":"2007","unstructured":"Koressaar, T. & Remm, M. Enhancements and modifications of primer design program Primer3. Bioinformatics 23, 1289\u20131291. https:\/\/doi.org\/10.1093\/bioinformatics\/btm091 (2007).","journal-title":"Bioinformatics"},{"key":"75243_CR15","doi-asserted-by":"publisher","first-page":"e115","DOI":"10.1093\/nar\/gks596","volume":"40","author":"A Untergasser","year":"2012","unstructured":"Untergasser, A. et al. Primer3\u2014New capabilities and interfaces. Nucleic Acids Res. 40, e115\u2013e115. https:\/\/doi.org\/10.1093\/nar\/gks596 (2012).","journal-title":"Nucleic Acids Res."},{"key":"75243_CR16","doi-asserted-by":"publisher","first-page":"3047","DOI":"10.1093\/bioinformatics\/btw354","volume":"32","author":"P Ewels","year":"2016","unstructured":"Ewels, P., Magnusson, M., Lundin, S. & K\u00e4ller, M. MultiQC: Summarize analysis results for multiple tools and samples in a single report. Bioinformatics 32, 3047\u20133048. https:\/\/doi.org\/10.1093\/bioinformatics\/btw354 (2016).","journal-title":"Bioinformatics"},{"key":"75243_CR17","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1093\/bioinformatics\/bts635","volume":"29","author":"A Dobin","year":"2013","unstructured":"Dobin, A. et al. STAR: Ultrafast universal RNA-seq aligner. Bioinformatics 29, 15\u201321. https:\/\/doi.org\/10.1093\/bioinformatics\/bts635 (2013).","journal-title":"Bioinformatics"},{"key":"75243_CR18","doi-asserted-by":"publisher","first-page":"644","DOI":"10.1038\/nbt.1883","volume":"29","author":"MG Grabherr","year":"2011","unstructured":"Grabherr, M. G. et al. Full-length transcriptome assembly from RNA-Seq data without a reference genome. Nat. Biotechnol. 29, 644\u2013652. https:\/\/doi.org\/10.1038\/nbt.1883 (2011).","journal-title":"Nat. Biotechnol."},{"key":"75243_CR19","doi-asserted-by":"publisher","first-page":"2184","DOI":"10.1093\/bioinformatics\/bts356","volume":"28","author":"L Wang","year":"2012","unstructured":"Wang, L., Wang, S. & Li, W. RSeQC: Quality control of RNA-seq experiments. Bioinformatics 28, 2184\u20132185. https:\/\/doi.org\/10.1093\/bioinformatics\/bts356 (2012).","journal-title":"Bioinformatics"},{"key":"75243_CR20","doi-asserted-by":"publisher","first-page":"178","DOI":"10.1093\/bib\/bbs017","volume":"14","author":"H Thorvaldsd\u00f3ttir","year":"2012","unstructured":"Thorvaldsd\u00f3ttir, H., Robinson, J. T. & Mesirov, J. P. Integrative Genomics Viewer (IGV): High-performance genomics data visualization and exploration. Brief. Bioinform. 14, 178\u2013192. https:\/\/doi.org\/10.1093\/bib\/bbs017 (2012).","journal-title":"Brief. Bioinform."},{"key":"75243_CR21","doi-asserted-by":"publisher","first-page":"e1006360","DOI":"10.1371\/journal.pcbi.1006360","volume":"14","author":"D Garrido-Mart\u00edn","year":"2018","unstructured":"Garrido-Mart\u00edn, D., Palumbo, E., Guig\u00f3, R. & Breschi, A. ggsashimi: Sashimi plot revised for browser- and annotation-independent splicing visualization. PLoS Comput. Biol. 14, e1006360. https:\/\/doi.org\/10.1371\/journal.pcbi.1006360 (2018).","journal-title":"PLoS Comput. Biol."},{"key":"75243_CR22","doi-asserted-by":"publisher","first-page":"252","DOI":"10.1016\/j.tvjl.2006.08.017","volume":"174","author":"AC Sewell","year":"2007","unstructured":"Sewell, A. C., Haskins, M. E. & Giger, U. Inherited metabolic disease in companion animals: Searching for nature\u2019s mistakes. Vet. J. 174, 252\u2013259. https:\/\/doi.org\/10.1016\/j.tvjl.2006.08.017 (2007).","journal-title":"Vet. J."},{"key":"75243_CR23","doi-asserted-by":"publisher","first-page":"122","DOI":"10.1093\/ilar.50.2.122","volume":"50","author":"DD Koeberl","year":"2009","unstructured":"Koeberl, D. D., Pinto, C., Brown, T. & Chen, Y. T. Gene therapy for inherited metabolic disorders in companion animals. ILAR J. 50, 122\u2013127. https:\/\/doi.org\/10.1093\/ilar.50.2.122 (2009).","journal-title":"ILAR J."},{"key":"75243_CR24","doi-asserted-by":"publisher","first-page":"143","DOI":"10.1111\/jsap.12039","volume":"54","author":"JRS Dandrieux","year":"2013","unstructured":"Dandrieux, J. R. S., Noble, P. J. M., Halladay, L. J., McLean, L. & German, A. J. Canine breed predispositions for marked hypocobalaminaemia or decreased folate concentration assessed by a laboratory survey. J. Small Anim. Pract. 54, 143\u2013148. https:\/\/doi.org\/10.1111\/jsap.12039 (2013).","journal-title":"J. Small Anim. Pract."},{"key":"75243_CR25","doi-asserted-by":"publisher","first-page":"104","DOI":"10.1053\/j.tcam.2012.04.004","volume":"27","author":"AJ German","year":"2012","unstructured":"German, A. J. Exocrine pancreatic insufficiency in the dog: Breed associations, nutritional considerations, and long-term outcome. Topics Companion Animal Med. 27, 104\u2013108. https:\/\/doi.org\/10.1053\/j.tcam.2012.04.004 (2012).","journal-title":"Topics Companion Animal Med."},{"key":"75243_CR26","doi-asserted-by":"publisher","first-page":"1404","DOI":"10.2741\/e470","volume":"4","author":"AE Jergens","year":"2012","unstructured":"Jergens, A. E. & Simpson, K. W. Inflammatory bowel disease in veterinary medicine. Front. Biosci. (Elite Edition). 4, 1404 (2012).","journal-title":"Front. Biosci. (Elite Edition)."},{"key":"75243_CR27","doi-asserted-by":"publisher","first-page":"210","DOI":"10.1016\/j.molmet.2016.01.001","volume":"5","author":"TA Bowman","year":"2016","unstructured":"Bowman, T. A. et al. Acyl CoA synthetase 5 (ACSL5) ablation in mice increases energy expenditure and insulin sensitivity and delays fat absorption. Mol. Metab. 5, 210\u2013220. https:\/\/doi.org\/10.1016\/j.molmet.2016.01.001 (2016).","journal-title":"Mol. Metab."},{"key":"75243_CR28","doi-asserted-by":"publisher","first-page":"2004","DOI":"10.1194\/jlr.M600150-JLR200","volume":"47","author":"DG Mashek","year":"2006","unstructured":"Mashek, D. G., Li, L. O. & Coleman, R. A. Rat long-chain acyl-CoA synthetase mRNA, protein, and activity vary in tissue distribution and in response to diet. J. Lipid Res. 47, 2004\u20132010. https:\/\/doi.org\/10.1194\/jlr.M600150-JLR200 (2006).","journal-title":"J. Lipid Res."},{"key":"75243_CR29","doi-asserted-by":"publisher","first-page":"945","DOI":"10.1074\/jbc.M507646200","volume":"281","author":"DG Mashek","year":"2006","unstructured":"Mashek, D. G., McKenzie, M. A., Van Horn, C. G. & Coleman, R. A. Rat long chain Acyl-CoA synthetase 5 increases fatty acid uptake and partitioning to cellular triacylglycerol in McArdle-RH7777 cells. J. Biol. Chem. 281, 945\u2013950. https:\/\/doi.org\/10.1074\/jbc.M507646200 (2006).","journal-title":"J. Biol. Chem."},{"key":"75243_CR30","doi-asserted-by":"publisher","first-page":"888","DOI":"10.1194\/jlr.M024562","volume":"53","author":"M Poppelreuther","year":"2012","unstructured":"Poppelreuther, M. et al. The N-terminal region of acyl-CoA synthetase 3 is essential for both the localization on lipid droplets and the function in fatty acid uptake. J. Lipid Res. 53, 888\u2013900. https:\/\/doi.org\/10.1194\/jlr.M024562 (2012).","journal-title":"J. Lipid Res."},{"key":"75243_CR31","doi-asserted-by":"publisher","first-page":"677","DOI":"10.1113\/JP272962","volume":"595","author":"BG Teodoro","year":"2017","unstructured":"Teodoro, B. G. et al. Long-chain acyl-CoA synthetase 6 regulates lipid synthesis and mitochondrial oxidative capacity in human and rat skeletal muscle. J. Physiol. 595, 677\u2013693. https:\/\/doi.org\/10.1113\/JP272962 (2017).","journal-title":"J. Physiol."},{"key":"75243_CR32","doi-asserted-by":"publisher","first-page":"655","DOI":"10.3892\/ijmm.2011.624","volume":"27","author":"M Wu","year":"2011","unstructured":"Wu, M., Cao, A., Dong, B. & Liu, J. Reduction of serum free fatty acids and triglycerides by liver-targeted expression of long chain acyl-CoA synthetase 3. Int. J. Mol. Med. 27, 655\u2013662. https:\/\/doi.org\/10.3892\/ijmm.2011.624 (2011).","journal-title":"Int. J. Mol. Med."},{"key":"75243_CR33","doi-asserted-by":"publisher","first-page":"1068","DOI":"10.1038\/oby.2007.630","volume":"15","author":"KB Adamo","year":"2007","unstructured":"Adamo, K. B. et al. Peroxisome proliferator-activated receptor \u03b3 2 and Acyl-CoA synthetase 5 polymorphisms influence diet response. Obesity 15, 1068\u20131075. https:\/\/doi.org\/10.1038\/oby.2007.630 (2007).","journal-title":"Obesity"},{"key":"75243_CR34","doi-asserted-by":"publisher","first-page":"53","DOI":"10.1016\/j.cmet.2010.05.012","volume":"12","author":"JM Ellis","year":"2010","unstructured":"Ellis, J. M. et al. Adipose acyl-CoA synthetase-1 directs fatty acids toward beta-oxidation and is required for cold thermogenesis. Cell Metab. 12, 53\u201364. https:\/\/doi.org\/10.1016\/j.cmet.2010.05.012 (2010).","journal-title":"Cell Metab."},{"key":"75243_CR35","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1016\/j.molmet.2018.01.012","volume":"9","author":"EA Killion","year":"2018","unstructured":"Killion, E. A. et al. A role for long-chain acyl-CoA synthetase-4 (ACSL4) in diet-induced phospholipid remodeling and obesity-associated adipocyte dysfunction. Mol. Metab. 9, 43\u201356. https:\/\/doi.org\/10.1016\/j.molmet.2018.01.012 (2018).","journal-title":"Mol. Metab."},{"key":"75243_CR36","doi-asserted-by":"publisher","first-page":"735","DOI":"10.1016\/S0026-0495(03)00034-9","volume":"52","author":"JD Privette","year":"2003","unstructured":"Privette, J. D., Hickner, R. C., MacDonald, K. G., Pories, W. J. & Barakat, H. A. Fatty acid oxidation by skeletal muscle homogenates from morbidly obese black and white American women. Metabolism 52, 735\u2013738. https:\/\/doi.org\/10.1016\/S0026-0495(03)00034-9 (2003).","journal-title":"Metabolism"},{"key":"75243_CR37","doi-asserted-by":"publisher","first-page":"56","DOI":"10.1186\/s12881-016-0320-4","volume":"17","author":"A Rajkumar","year":"2016","unstructured":"Rajkumar, A. et al. Acyl-CoA synthetase long-chain 5 genotype is associated with body composition changes in response to lifestyle interventions in postmenopausal women with overweight and obesity: A genetic association study on cohorts Montr\u00e9al-Ottawa New Emerging Team, and Complications Associated with Obesity. BMC Med. Genet. 17, 56. https:\/\/doi.org\/10.1186\/s12881-016-0320-4 (2016).","journal-title":"BMC Med. Genet."},{"key":"75243_CR38","doi-asserted-by":"publisher","first-page":"271","DOI":"10.1016\/j.metabol.2018.03.019","volume":"83","author":"A Rajkumar","year":"2018","unstructured":"Rajkumar, A. et al. ACSL5 genotype influence on fatty acid metabolism: A cellular, tissue, and whole-body study. Metabolism 83, 271\u2013279. https:\/\/doi.org\/10.1016\/j.metabol.2018.03.019 (2018).","journal-title":"Metabolism"},{"key":"75243_CR39","doi-asserted-by":"publisher","first-page":"141","DOI":"10.1007\/s10709-019-00057-3","volume":"147","author":"C Zhao","year":"2019","unstructured":"Zhao, C. et al. Adaptive evolution of the ACSL gene family in Carnivora. Genetica 147, 141\u2013148. https:\/\/doi.org\/10.1007\/s10709-019-00057-3 (2019).","journal-title":"Genetica"},{"key":"75243_CR40","doi-asserted-by":"publisher","first-page":"e1006661","DOI":"10.1371\/journal.pgen.1006661","volume":"13","author":"J Plassais","year":"2017","unstructured":"Plassais, J. et al. Analysis of large versus small dogs reveals three genes on the canine X chromosome associated with body weight, muscling and back fat thickness. PLoS Genet. 13, e1006661. https:\/\/doi.org\/10.1371\/journal.pgen.1006661 (2017).","journal-title":"PLoS Genet."},{"key":"75243_CR41","doi-asserted-by":"publisher","first-page":"88","DOI":"10.1186\/1476-511X-12-88","volume":"12","author":"N Meller","year":"2013","unstructured":"Meller, N., Morgan, M. E., Wong, W. P., Altemus, J. B. & Sehayek, E. Targeting of Acyl-CoA synthetase 5 decreases jejunal fatty acid activation with no effect on dietary long-chain fatty acid absorption. Lipids Health Dis. 12, 88\u201388. https:\/\/doi.org\/10.1186\/1476-511X-12-88 (2013).","journal-title":"Lipids Health Dis."},{"key":"75243_CR42","doi-asserted-by":"publisher","first-page":"309","DOI":"10.1038\/326309a0","volume":"326","author":"BS Glick","year":"1987","unstructured":"Glick, B. S. & Rothman, J. E. Possible role for fatty acyl-coenzyme A in intracellular protein transport. Nature 326, 309\u2013312. https:\/\/doi.org\/10.1038\/326309a0 (1987).","journal-title":"Nature"},{"key":"75243_CR43","doi-asserted-by":"publisher","first-page":"2880","DOI":"10.1073\/pnas.94.7.2880","volume":"94","author":"M-J Kang","year":"1997","unstructured":"Kang, M.-J. et al. A novel arachidonate-preferring acyl-CoA synthetase is present in steroidogenic cells of the rat adrenal, ovary, and testis. Proc. Natl. Acad. Sci. 94, 2880. https:\/\/doi.org\/10.1073\/pnas.94.7.2880 (1997).","journal-title":"Proc. Natl. Acad. Sci."},{"key":"75243_CR44","doi-asserted-by":"publisher","first-page":"679","DOI":"10.1093\/oxfordjournals.jbchem.a022165","volume":"124","author":"E Oikawa","year":"1998","unstructured":"Oikawa, E. et al. A novel Acyl-CoA synthetase, ACS5, expressed in intestinal epithelial cells and proliferating preadipocytes1. J. Biochem. 124, 679\u2013685. https:\/\/doi.org\/10.1093\/oxfordjournals.jbchem.a022165 (1998).","journal-title":"J. Biochem."},{"key":"75243_CR45","doi-asserted-by":"publisher","first-page":"188","DOI":"10.1002\/path.1504","volume":"202","author":"N Gassler","year":"2004","unstructured":"Gassler, N. et al. Expression of acyl-CoA synthetase 5 reflects the state of villus architecture in human small intestine. J. Pathol. 202, 188\u2013196. https:\/\/doi.org\/10.1002\/path.1504 (2004).","journal-title":"J. Pathol."},{"key":"75243_CR46","doi-asserted-by":"publisher","first-page":"587","DOI":"10.1053\/j.gastro.2007.06.005","volume":"133","author":"N Gassler","year":"2007","unstructured":"Gassler, N. et al. Regulation of enterocyte apoptosis by Acyl-CoA synthetase 5 splicing. Gastroenterology 133, 587\u2013598. https:\/\/doi.org\/10.1053\/j.gastro.2007.06.005 (2007).","journal-title":"Gastroenterology"},{"key":"75243_CR47","doi-asserted-by":"publisher","first-page":"1084","DOI":"10.1136\/gut.2005.073262","volume":"55","author":"N Gassler","year":"2006","unstructured":"Gassler, N. et al. Molecular characterisation of non-absorptive and absorptive enterocytes in human small intestine. Gut 55, 1084\u20131089. https:\/\/doi.org\/10.1136\/gut.2005.073262 (2006).","journal-title":"Gut"},{"key":"75243_CR48","doi-asserted-by":"publisher","first-page":"3270","DOI":"10.1194\/jlr.M009407","volume":"51","author":"SY Bu","year":"2010","unstructured":"Bu, S. Y. & Mashek, D. G. Hepatic long-chain acyl-CoA synthetase 5 mediates fatty acid channeling between anabolic and catabolic pathways. J. Lipid Res. 51, 3270\u20133280. https:\/\/doi.org\/10.1194\/jlr.M009407 (2010).","journal-title":"J. Lipid Res."},{"key":"75243_CR49","doi-asserted-by":"publisher","first-page":"2766","DOI":"10.1111\/febs.12263","volume":"280","author":"S Blaskovic","year":"2013","unstructured":"Blaskovic, S., Blanc, M. & van der Goot, F. G. What does S-palmitoylation do to membrane proteins?. FEBS J. 280, 2766\u20132774. https:\/\/doi.org\/10.1111\/febs.12263 (2013).","journal-title":"FEBS J."},{"key":"75243_CR50","doi-asserted-by":"publisher","DOI":"10.3390\/ijms19123820","author":"M Spinelli","year":"2018","unstructured":"Spinelli, M., Fusco, S. & Grassi, C. Nutrient-dependent changes of protein palmitoylation: Impact on nuclear enzymes and regulation of gene expression. Int. J. Mol. Sci. https:\/\/doi.org\/10.3390\/ijms19123820 (2018).","journal-title":"Int. J. Mol. Sci."},{"key":"75243_CR51","doi-asserted-by":"publisher","DOI":"10.1101\/431106","author":"PA Sandoz","year":"2018","unstructured":"Sandoz, P. A. et al. The architecture of the endoplasmic reticulum is regulated by the reversible lipid modification of the shaping protein CLIMP-63. bioRxiv. https:\/\/doi.org\/10.1101\/431106 (2018).","journal-title":"bioRxiv."},{"key":"75243_CR52","doi-asserted-by":"publisher","first-page":"e27826","DOI":"10.7554\/eLife.27826","volume":"6","author":"L Abrami","year":"2017","unstructured":"Abrami, L. et al. Identification and dynamics of the human ZDHHC16-ZDHHC6 palmitoylation cascade. Elife 6, e27826. https:\/\/doi.org\/10.7554\/eLife.27826 (2017).","journal-title":"Elife"},{"key":"75243_CR53","doi-asserted-by":"publisher","first-page":"2488","DOI":"10.1016\/j.febslet.2012.06.011","volume":"586","author":"M Fairbank","year":"2012","unstructured":"Fairbank, M., Huang, K., El-Husseini, A. & Nabi, I. R. RING finger palmitoylation of the endoplasmic reticulum Gp78 E3 ubiquitin ligase. FEBS Lett. 586, 2488\u20132493. https:\/\/doi.org\/10.1016\/j.febslet.2012.06.011 (2012).","journal-title":"FEBS Lett."},{"key":"75243_CR54","doi-asserted-by":"publisher","first-page":"16478","DOI":"10.1073\/pnas.1417176111","volume":"111","author":"GJ Fredericks","year":"2014","unstructured":"Fredericks, G. J. et al. Stable expression and function of the inositol 1,4,5-triphosphate receptor requires palmitoylation by a DHHC6\/selenoprotein K complex. Proc. Natl. Acad. Sci. 111, 16478. https:\/\/doi.org\/10.1073\/pnas.1417176111 (2014).","journal-title":"Proc. Natl. Acad. Sci."},{"key":"75243_CR55","doi-asserted-by":"publisher","first-page":"1823","DOI":"10.1038\/emboj.2012.15","volume":"31","author":"AKK Lakkaraju","year":"2012","unstructured":"Lakkaraju, A. K. K. et al. Palmitoylated calnexin is a key component of the ribosome\u2013translocon complex. EMBO J. 31, 1823\u20131835. https:\/\/doi.org\/10.1038\/emboj.2012.15 (2012).","journal-title":"EMBO J."},{"key":"75243_CR56","doi-asserted-by":"crossref","unstructured":"Senyilmaz, D. et al. Regulation of mitochondrial morphology and function by stearoylation of TFR1. Nature 525, 124. 10.1038\/nature14601. https:\/\/www.nature.com\/articles\/nature14601#supplementary-information (2015).","DOI":"10.1038\/nature14601"},{"key":"75243_CR57","doi-asserted-by":"publisher","first-page":"261","DOI":"10.12688\/f1000research.6464.1","volume":"4","author":"M Blanc","year":"2015","unstructured":"Blanc, M. et al. SwissPalm: Protein palmitoylation database. F1000Res 4, 261\u2013261. https:\/\/doi.org\/10.12688\/f1000research.6464.1 (2015).","journal-title":"F1000Res"},{"key":"75243_CR58","doi-asserted-by":"publisher","first-page":"e101314","DOI":"10.1371\/journal.pone.0101314","volume":"9","author":"H-C Lo","year":"2014","unstructured":"Lo, H.-C., Yang, R.-B. & Lee, C.-H. Guanylyl cyclase-G modulates Jejunal apoptosis and inflammation in mice with intestinal ischemia and reperfusion. PLoS ONE 9, e101314. https:\/\/doi.org\/10.1371\/journal.pone.0101314 (2014).","journal-title":"PLoS ONE"},{"key":"75243_CR59","doi-asserted-by":"publisher","first-page":"e884","DOI":"10.1371\/journal.pone.0000884","volume":"2","author":"JM Young","year":"2007","unstructured":"Young, J. M., Waters, H., Dong, C., F\u00fclle, H.-J. & Liman, E. R. Degeneration of the olfactory guanylyl cyclase D Gene during PRIMATE evolution. PLoS ONE 2, e884. https:\/\/doi.org\/10.1371\/journal.pone.0000884 (2007).","journal-title":"PLoS ONE"},{"key":"75243_CR60","doi-asserted-by":"publisher","first-page":"e91172","DOI":"10.1371\/journal.pone.0091172","volume":"9","author":"MP Hoeppner","year":"2014","unstructured":"Hoeppner, M. P. et al. An improved canine genome and a comprehensive catalogue of coding genes and non-coding transcripts. PLoS ONE 9, e91172. https:\/\/doi.org\/10.1371\/journal.pone.0091172 (2014).","journal-title":"PLoS ONE"},{"key":"75243_CR61","doi-asserted-by":"publisher","first-page":"132","DOI":"10.1007\/s40142-017-0125-6","volume":"5","author":"ES Goetzman","year":"2017","unstructured":"Goetzman, E. S. Advances in the understanding and treatment of mitochondrial fatty acid oxidation disorders. Curr. Genet. Med. Rep. 5, 132\u2013142. https:\/\/doi.org\/10.1007\/s40142-017-0125-6 (2017).","journal-title":"Curr. Genet. Med. Rep."},{"key":"75243_CR62","doi-asserted-by":"publisher","first-page":"555","DOI":"10.1007\/s10545-010-9188-1","volume":"33","author":"U Spiekerkoetter","year":"2010","unstructured":"Spiekerkoetter, U. et al. Current issues regarding treatment of mitochondrial fatty acid oxidation disorders. J. Inherit. Metab. Dis. 33, 555\u2013561. https:\/\/doi.org\/10.1007\/s10545-010-9188-1 (2010).","journal-title":"J. Inherit. Metab. Dis."},{"key":"75243_CR63","doi-asserted-by":"publisher","first-page":"110","DOI":"10.1016\/j.ymgme.2011.09.030","volume":"105","author":"AM Behrend","year":"2012","unstructured":"Behrend, A. M. et al. Substrate oxidation and cardiac performance during exercise in disorders of long chain fatty acid oxidation. Mol. Genet. Metab. 105, 110\u2013115. https:\/\/doi.org\/10.1016\/j.ymgme.2011.09.030 (2012).","journal-title":"Mol. Genet. Metab."},{"key":"75243_CR64","doi-asserted-by":"publisher","first-page":"58","DOI":"10.1016\/j.ymgme.2006.06.004","volume":"89","author":"MB Gillingham","year":"2006","unstructured":"Gillingham, M. B., Scott, B., Elliott, D. & Harding, C. O. Metabolic control during exercise with and without medium-chain triglycerides (MCT) in children with long-chain 3-hydroxy acyl-CoA dehydrogenase (LCHAD) or trifunctional protein (TFP) deficiency. Mol. Genet. Metab. 89, 58\u201363. https:\/\/doi.org\/10.1016\/j.ymgme.2006.06.004 (2006).","journal-title":"Mol. Genet. Metab."},{"key":"75243_CR65","doi-asserted-by":"publisher","first-page":"95","DOI":"10.1007\/8904_2016_561","volume-title":"JIMD Reports, Vol. 32","author":"DCD van Eerd","year":"2017","unstructured":"van Eerd, D. C. D. et al. Management of an LCHADD patient during pregnancy and high intensity exercise. In JIMD Reports, Vol. 32, 95\u2013100. https:\/\/doi.org\/10.1007\/8904_2016_561 (Springer, Berlin, Heidelberg, 2017)."},{"key":"75243_CR66","doi-asserted-by":"publisher","first-page":"306","DOI":"10.1046\/j.1439-0396.2002.00387.x","volume":"86","author":"AC Beynen","year":"2002","unstructured":"Beynen, A. C., Kappert, H. J., Lemmens, A. G. & Van Dongen, A. M. Plasma lipid concentrations, macronutrient digestibility and mineral absorption in dogs fed a dry food containing medium-chain triglycerides. J. Animal Physiol. Animal Nutr. 86, 306\u2013312. https:\/\/doi.org\/10.1046\/j.1439-0396.2002.00387.x (2002).","journal-title":"J. Animal Physiol. Animal Nutr."},{"key":"75243_CR67","doi-asserted-by":"publisher","first-page":"35","DOI":"10.1016\/j.fct.2008.06.080","volume":"47","author":"RA Matulka","year":"2009","unstructured":"Matulka, R. A., Thompson, D. V. M. L. & Burdock, G. A. Lack of toxicity by medium chain triglycerides (MCT) in canines during a 90-day feeding study. Food Chem. Toxicol. 47, 35\u201339. https:\/\/doi.org\/10.1016\/j.fct.2008.06.080 (2009).","journal-title":"Food Chem. Toxicol."},{"key":"75243_CR68","doi-asserted-by":"publisher","first-page":"1293","DOI":"10.2460\/ajvr.2004.65.1293","volume":"65","author":"GM Rutz","year":"2004","unstructured":"Rutz, G. M., Steiner, J. M., Bauer, J. E. & Williams, D. A. Effects of exchange of dietary medium chain triglycerides for long-chain triglycerides on serum biochemical variables and subjectively assessed well-being of dogs with exocrine pancreatic insufficiency. Am. J. Vet. Res. 65, 1293. https:\/\/doi.org\/10.2460\/ajvr.2004.65.1293 (2004).","journal-title":"Am. J. Vet. Res."},{"key":"75243_CR69","doi-asserted-by":"publisher","first-page":"927","DOI":"10.1093\/ajcn\/45.5.927","volume":"45","author":"R Cotter","year":"1987","unstructured":"Cotter, R., Taylor, C. A., Johnson, R. & Rowe, W. B. A metabolic comparison of a pure long-chain triglyceride lipid emulsion (LCT) and various medium-chain triglyceride (MCT)-LCT combination emulsions in dogs. Am. J. Clin. Nutr. 45, 927\u2013939. https:\/\/doi.org\/10.1093\/ajcn\/45.5.927 (1987).","journal-title":"Am. J. Clin. Nutr."}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-020-75243-x.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-020-75243-x","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-020-75243-x.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,12,6]],"date-time":"2022-12-06T21:58:01Z","timestamp":1670363881000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-020-75243-x"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,10,26]]},"references-count":69,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2020,12]]}},"alternative-id":["75243"],"URL":"https:\/\/doi.org\/10.1038\/s41598-020-75243-x","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2020,10,26]]},"assertion":[{"value":"12 June 2020","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"6 October 2020","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"26 October 2020","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":"18223"}}