{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,14]],"date-time":"2026-04-14T03:49:13Z","timestamp":1776138553026,"version":"3.50.1"},"reference-count":85,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,1,27]],"date-time":"2026-01-27T00:00:00Z","timestamp":1769472000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Endocrinol."],"abstract":"<jats:p>\n                    Appetite regulation in fish relies on complex neuroendocrine pathways within the brain-gut axis, with ghrelin (Ghrl) and cholecystokinin (Cck) as central players. However, their spatial distribution along the gastrointestinal tract (GIT) and responses to feeding status remain poorly understood in teleosts. This study investigated: i) the baseline distribution of Ghrl and Cck levels along the GIT of juvenile\n                    <jats:italic>Sparus aurata<\/jats:italic>\n                    fed a commercial diet; ii) their temporal dynamics during short-term fasting and refeeding; and iii) the influence of diet composition on their spatiotemporal profiles. Juveniles were fed for 92 days with: i) a control diet containing 20% fishmeal (CT); ii) a plant protein diet replacing 60% of fishmeal with hydrolyzed plant protein (PP); and iii) the PP diet supplemented with 2%\n                    <jats:italic>\n                      LB-Green\n                      <jats:sub>Grape<\/jats:sub>\n                    <\/jats:italic>\n                    functional additive (GG). Fish were sampled at 2, 6, and 24\u00a0h post-feeding (Cf), after 7 days of fasting (Ft), and at 2, 6, and 24\u00a0h post-refeeding (Rf). Hormone levels were quantified across five GIT segments, including the stomach (S1) and four equal intestinal segments (S2\u2013S5). Baseline characterization revealed elevated Ghrl content in S3 and S5, whereas Cck levels were highest in S5. During fasting, Ghrl levels declined, while Cck increased in S1, S2, and S5 with distinct temporal patterns. After refeeding, gastric Ghrl levels (S1) decreased within 24\u00a0h, potentially reflecting secretion into plasma and involvement in hunger signaling, although plasma levels were not measured. In contrast, Cck levels in the anterior intestine (S2) rose sharply 24\u00a0h after refeeding, suggesting an anticipatory response to refeeding, possibly related to a dual role involving both rapid satiety signaling and preparatory modulation of digestive activity. The PP and GG diets maintained high gastric Ghrl (S1) and lowered intestinal Cck (S2) levels after feeding, especially in the PP diet. This pattern may either prolong satiety and reduce feed intake or reflect changes in hormone release due to lower caloric intake, with the PP diet lowering growth and feed efficiency, partially offset by the functional additive. The study maps Ghrl and Cck in the\n                    <jats:italic>S. aurata<\/jats:italic>\n                    GIT, showing spatial, temporal, and dietary regulation, with implications for aquaculture nutrition.\n                  <\/jats:p>","DOI":"10.3389\/fendo.2026.1734169","type":"journal-article","created":{"date-parts":[[2026,1,27]],"date-time":"2026-01-27T05:25:22Z","timestamp":1769491522000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":1,"title":["Spatiotemporal characterization of ghrelin and cholecystokinin levels in the gastrointestinal tract of juvenile Sparus aurata: effects of feeding status and diet composition"],"prefix":"10.3389","volume":"17","author":[{"given":"Anyell","family":"Caderno","sequence":"first","affiliation":[{"name":"Department of Biology, Faculty of Marine and Environmental Sciences, Instituto Universitario de Investigaci\u00f3n Marina (INMAR), University of C\u00e1diz","place":["Puerto Real, Spain"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Patrik","family":"Tang","sequence":"additional","affiliation":[{"name":"NORCE Research AS, Climate & Environment Department, Fish Biology and Aquaculture (FBA) Group","place":["Bergen, Norway"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ver\u00f3nica","family":"de las Heras","sequence":"additional","affiliation":[{"name":"Department of Biology, Faculty of Marine and Environmental Sciences, Instituto Universitario de Investigaci\u00f3n Marina (INMAR), University of C\u00e1diz","place":["Puerto Real, Spain"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Naouel","family":"Gharbi","sequence":"additional","affiliation":[{"name":"NORCE Research AS, Climate & Environment Department, Fish Biology and Aquaculture (FBA) Group","place":["Bergen, Norway"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Francisco Javier","family":"Alarc\u00f3n-L\u00f3pez","sequence":"additional","affiliation":[{"name":"Department of Biology and Geology, Faculty of Experimental Sciences, University of Almer\u00eda","place":["Almer\u00eda, Spain"]},{"name":"LifeBioencapsulation S.L., Parque Cient\u00edfico PITA","place":["Almer\u00eda, Spain"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Juan Miguel","family":"Mancera","sequence":"additional","affiliation":[{"name":"Department of Biology, Faculty of Marine and Environmental Sciences, Instituto Universitario de Investigaci\u00f3n Marina (INMAR), University of C\u00e1diz","place":["Puerto Real, Spain"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Juan Antonio","family":"Martos-Sitcha","sequence":"additional","affiliation":[{"name":"Department of Biology, Faculty of Marine and Environmental Sciences, Instituto Universitario de Investigaci\u00f3n Marina (INMAR), University of C\u00e1diz","place":["Puerto Real, Spain"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Neda","family":"Gilannejad","sequence":"additional","affiliation":[{"name":"NORCE Research AS, Climate & Environment Department, Fish Biology and Aquaculture (FBA) Group","place":["Bergen, Norway"]}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1965","published-online":{"date-parts":[[2026,1,27]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","DOI":"10.3389\/fendo.2021.798903","article-title":"Fish feed intake, feeding behavior, and the physiological response of apelin to fasting and refeeding","volume":"12","author":"Assan","year":"2021","journal-title":"Front Endocrinol"},{"key":"B2","doi-asserted-by":"publisher","DOI":"10.1016\/j.ygcen.2011.12.025","article-title":"The comparative endocrinology of feeding in fish: Insights and challenges","volume":"176","author":"Hoskins","year":"2012","journal-title":"Gen Comp Endocrinol"},{"key":"B3","doi-asserted-by":"publisher","first-page":"79","DOI":"10.1016\/j.ygcen.2013.03.013","article-title":"The role of ghrelin in energy balance regulation in fish","volume":"187","author":"J\u00f6nsson","year":"2013","journal-title":"Gen Comp Endocrinol"},{"key":"B4","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1002\/9781444354614.ch7","article-title":"Appetite and Feed Intake","volume-title":"Aquaculture and Behavior","author":"Jobling","year":"2012"},{"key":"B5","doi-asserted-by":"publisher","first-page":"8","DOI":"10.1016\/j.cbpa.2008.12.001","article-title":"Aspects of the hormonal regulation of appetite in fish with emphasis on goldfish, Atlantic cod and winter flounder: Notes on actions and responses to nutritional, environmental and reproductive changes","volume":"153","author":"Volkoff","year":"2009","journal-title":"Comp Biochem Physiol A Mol Integr Physiol"},{"key":"B6","doi-asserted-by":"publisher","DOI":"10.1016\/j.cbpb.2004.09.007","article-title":"Hormonal regulation of the fish gastrointestinal tract","volume":"139","author":"Buddington","year":"2004","journal-title":"Comp Biochem Physiol A Mol Integr Physiol"},{"key":"B7","doi-asserted-by":"publisher","DOI":"10.1016\/j.ygcen.2006.01.011","article-title":"Gastroenteropancreatic hormones and metabolism in fish","volume":"148","author":"Nelson","year":"2006","journal-title":"Gen Comp Endocrinol"},{"key":"B8","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1172\/JCI30227","article-title":"Gastrointestinal regulation of food intake","volume":"117","author":"Cummings","year":"2007","journal-title":"J Clin Invest"},{"key":"B9","doi-asserted-by":"publisher","DOI":"10.1530\/JME-17-0320","article-title":"Central regulation of food intake in fish: an evolutionary perspective","volume":"60","author":"Soengas","year":"2018","journal-title":"J Mol Endocrinol"},{"key":"B10","doi-asserted-by":"publisher","DOI":"10.3389\/fendo.2017.00073","article-title":"Appetite-controlling endocrine systems in teleosts","volume":"8","author":"R\u00f8nnestad","year":"2017","journal-title":"Front Endocrinol"},{"key":"B11","doi-asserted-by":"publisher","DOI":"10.3389\/fnins.2016.00540","article-title":"The neuroendocrine regulation of food intake in fish: A review of current knowledge","volume":"10","author":"Volkoff","year":"2016","journal-title":"Front Neurosci"},{"key":"B12","doi-asserted-by":"publisher","first-page":"396","DOI":"10.1016\/j.cbpb.2005.02.011","article-title":"Structure, distribution and physiological functions of ghrelin in fish","volume":"140","author":"Unniappan","year":"2005","journal-title":"Comp Biochem Physiol A Mol Integr Physiol"},{"key":"B13","doi-asserted-by":"publisher","DOI":"10.1016\/j.peptides.2011.05.006","article-title":"Central and peripheral effects of ghrelin on energy balance, food intake and lipid metabolism in teleost fish","volume":"32","author":"Kang","year":"2011","journal-title":"Peptides"},{"key":"B14","doi-asserted-by":"publisher","DOI":"10.1016\/j.cbpa.2005.10.026","article-title":"The role of neuropeptide Y, orexins, cocaine and amphetamine-related transcript, cholecystokinin, amylin and leptin in the regulation of feeding in fish","volume":"144","author":"Volkoff","year":"2006","journal-title":"Comp Biochem Physiol A Mol Integr Physiol"},{"key":"B15","doi-asserted-by":"publisher","DOI":"10.1016\/j.ygcen.2008.11.002","article-title":"Fasting induces preproghrelin mRNA expression in the brain and gut of zebrafish, Danio rerio","volume":"161","author":"Amole","year":"2009","journal-title":"Gen Comp Endocrinol"},{"key":"B16","doi-asserted-by":"publisher","first-page":"e52428","DOI":"10.1371\/journal.pone.0052428","article-title":"Cholecystokinin in white sea bream: molecular cloning, regional expression, and immunohistochemical localization in the gut after feeding and fasting","volume":"7","author":"Micale","year":"2012","journal-title":"PloS One"},{"key":"B17","doi-asserted-by":"publisher","DOI":"10.3390\/ani12233287","article-title":"The interplay between central and peripheral systems in feed intake regulation in european seabass (Dicentrarchus labrax) juveniles","volume":"12","author":"Martins","year":"2022","journal-title":"Animals"},{"key":"B18","doi-asserted-by":"publisher","DOI":"10.3390\/biology13080635","article-title":"Role of cholecystokinin (cck) in feeding regulation of largemouth bass (Micropterus salmoides): peptide activation and antagonist inhibition","volume":"13","author":"Liang","year":"2024","journal-title":"Biology"},{"key":"B19","doi-asserted-by":"publisher","DOI":"10.1111\/anu.13138","article-title":"A feed is still only as good as its ingredients: An update on the nutritional research strategies for the optimal evaluation of ingredients for aquaculture feeds","volume":"26","author":"Glencross","year":"2020","journal-title":"Aquaculture Nutrition"},{"key":"B20","doi-asserted-by":"publisher","DOI":"10.1007\/s12562-012-0537-x","article-title":"Response of cholecystokinin and digestive enzyme mRNA levels to various feed ingredients in yellowtail Seriola quinqueradiata","volume":"78","author":"Furutani","year":"2012","journal-title":"Fish Sci"},{"key":"B21","doi-asserted-by":"publisher","DOI":"10.1016\/j.aquaculture.2021.737088","article-title":"Central regulation of food intake is not affected by inclusion of defatted Tenebrio molitor larvae meal in diets for European sea bass (Dicentrarchus labrax)","volume":"544","author":"Basto","year":"2021","journal-title":"Aquaculture"},{"key":"B22","doi-asserted-by":"publisher","DOI":"10.1016\/j.aquaculture.2022.738604","article-title":"Conde-Sieira M. Partial and total fishmeal replacement by defatted Tenebrio molitor larvae meal do not alter short- and mid-term regulation of food intake in European sea bass (Dicentrarchus labrax)","volume":"560","author":"Basto","year":"2022","journal-title":"Aquaculture"},{"key":"B23","doi-asserted-by":"publisher","DOI":"10.1016\/j.aquaculture.2020.735867","article-title":"The digestive function of gilthead seabream juveniles in relation to feeding frequency","volume":"531","author":"Gilannejad","year":"2021","journal-title":"Aquaculture"},{"key":"B24","doi-asserted-by":"publisher","DOI":"10.3389\/fendo.2018.00399","article-title":"Ghrelin and its receptors in gilthead sea bream: nutritional regulation","volume":"9","author":"Perell\u00f3-Amor\u00f3s","year":"2018","journal-title":"Front Endocrinol"},{"key":"B25","doi-asserted-by":"publisher","DOI":"10.1016\/j.ygcen.2016.10.003","article-title":"Effect of dietary macronutrients on the expression of cholecystokinin, leptin, ghrelin and neuropeptide Y in gilthead sea bream (Sparus aurata)","volume":"240","author":"Babaei","year":"2017","journal-title":"Gen Comp Endocrinol"},{"key":"B26","doi-asserted-by":"publisher","DOI":"10.3390\/ani11071919","article-title":"Appetite regulation, growth performances and fish quality are modulated by alternative dietary protein ingredients in gilthead sea bream (Sparus aurata) culture","volume":"11","author":"Pulido-Rodriguez","year":"2021","journal-title":"Animals"},{"key":"B27","doi-asserted-by":"publisher","DOI":"10.1016\/j.cbpa.2020.110868","article-title":"Daily rhythms of intestinal cholecystokinin and pancreatic proteases activity in Senegalese sole juveniles with diurnal and nocturnal feeding","volume":"253","author":"Gilannejad","year":"2021","journal-title":"Comp Biochem Physiol A Mol Integr Physiol"},{"key":"B28","doi-asserted-by":"publisher","first-page":"79","DOI":"10.1016\/0044-8486(94)00175-N","article-title":"Histology and histochemistry of the development of the digestive system of larval gilthead seabream, Sparus aurata L","volume":"130","author":"Sarasquete","year":"1995","journal-title":"Aquaculture"},{"key":"B29","doi-asserted-by":"publisher","first-page":"190","DOI":"10.1016\/0016-6480(87)90267-X","article-title":"Gastrin\/CCK-like immunoreactivity in endocrine cells and nerves in the gastrointestinal tract of the cod, Gadus morhua, and the effect of peptides of the gastrin\/CCK family on cod gastrointestinal smooth muscle","volume":"66","author":"J\u00f6nsson","year":"1987","journal-title":"Gen Comp Endocrinol"},{"key":"B30","doi-asserted-by":"publisher","first-page":"e20181165","DOI":"10.1590\/0001-3765202020181165","article-title":"Localization and distribution of CCK-8, NPY, Leu-ENK-, and Ghrelin- in the digestive tract of Prochilodus lineatus (Valenciennes, 1836)","volume":"92","author":"Barrios","year":"2020","journal-title":"Acad Bras Ci\u00eanc"},{"key":"B31","doi-asserted-by":"publisher","DOI":"10.1016\/j.repbre.2021.07.004","article-title":"A review on ghrelin and fish reproduction","volume":"1","author":"Zhong","year":"2021","journal-title":"Reprod Breed"},{"key":"B32","doi-asserted-by":"publisher","DOI":"10.1155\/2010\/945056","article-title":"Ghrelin cells in the gastrointestinal tract","volume":"2010","author":"Sakata","year":"2010","journal-title":"Int J Pept"},{"key":"B33","doi-asserted-by":"publisher","DOI":"10.1016\/j.peptides.2011.04.027","article-title":"Recent advances in the phylogenetic study of ghrelin","volume":"32","author":"Kaiya","year":"2011","journal-title":"Peptides"},{"key":"B34","doi-asserted-by":"publisher","DOI":"10.1210\/en.2002-220644","article-title":"Goldfish ghrelin: molecular characterization of the complementary deoxyribonucleic acid, partial gene structure and evidence for its stimulatory role in food intake","volume":"143","author":"Unniappan","year":"2002","journal-title":"Endocrinology"},{"key":"B35","doi-asserted-by":"publisher","DOI":"10.1210\/en.2003-1085","article-title":"Peptide purification, complementary deoxyribonucleic acid (DNA) and genomic DNA cloning, and functional characterization of ghrelin in rainbow trout","volume":"144","author":"Kaiya","year":"2003","journal-title":"Endocrinology"},{"key":"B36","doi-asserted-by":"publisher","DOI":"10.1016\/j.ygcen.2007.05.028","article-title":"Sea bass ghrelin: Molecular cloning and mRNA quantification during fasting and refeeding","volume":"155","author":"Terova","year":"2008","journal-title":"Gen Comp Endocrinol"},{"key":"B37","doi-asserted-by":"publisher","DOI":"10.1016\/j.ygcen.2015.05.004","article-title":"Cloning, tissue distribution and effects of fasting on mRNA expression levels of leptin and ghrelin in red-bellied piranha (Pygocentrus nattereri)","volume":"217","author":"Volkoff","year":"2015","journal-title":"Gen Comp Endocrinol"},{"key":"B38","doi-asserted-by":"publisher","DOI":"10.1016\/j.ygcen.2015.08.009","article-title":"Ghrelin, neuropeptide Y (NPY) and cholecystokinin (CCK) in blunt snout bream (Megalobrama amblycephala): cDNA cloning, tissue distribution and mRNA expression changes responding to fasting and refeeding","volume":"223","author":"Ji","year":"2015","journal-title":"Gen Comp Endocrinol"},{"key":"B39","doi-asserted-by":"publisher","DOI":"10.1002\/jemt.20709","article-title":"Distribution of ghrelin peptide in the gastrointestinal tract of stomachless and stomach-containing teleosts","volume":"72","author":"Arcamone","year":"2009","journal-title":"Microsc Res Tech"},{"key":"B40","doi-asserted-by":"publisher","DOI":"10.1016\/j.ygcen.2025.114824","article-title":"Does motilin primarily regulate gastrointestinal motility in fish","volume":"373","author":"Kitazawa","year":"2025","journal-title":"Gen Comp Endocrinol"},{"key":"B41","doi-asserted-by":"publisher","DOI":"10.1016\/j.ygcen.2007.05.016","article-title":"Identification of genes for the ghrelin and motilin receptors and a novel related gene in fish, and stimulation of intestinal motility in zebrafish (Danio rerio) by ghrelin and motilin","volume":"155","author":"Olsson","year":"2008","journal-title":"Gen Comp Endocrinol"},{"key":"B42","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.2006.116160","article-title":"Evidence that stimulation of ghrelin receptors in the spinal cord initiates propulsive activity in the colon of the rat","volume":"576","author":"Shimizu","year":"2006","journal-title":"J Physiol"},{"key":"B43","doi-asserted-by":"publisher","DOI":"10.1242\/jeb.202.2.161","article-title":"Cholecystokinin affects gastric emptying and stomach motility in the rainbow trout Oncorhynchus mykiss","volume":"202","author":"Olsson","year":"1999","journal-title":"J Exp Biol"},{"key":"B44","doi-asserted-by":"publisher","DOI":"10.3748\/wjg.v19.i12.1919","article-title":"Immunohistochemical study of the digestive tract of Oligosarcus hepsetus","volume":"19","author":"Vieira-Lopes","year":"2013","journal-title":"World J Gastroenterol WJG"},{"key":"B45","doi-asserted-by":"publisher","DOI":"10.12717\/DR.2016.20.3.207","article-title":"Digestive physiological characteristics of the gobiidae","volume":"20","author":"Hur","year":"2016","journal-title":"Dev Reprod"},{"key":"B46","doi-asserted-by":"publisher","DOI":"10.1016\/j.ygcen.2014.02.019","article-title":"Cholecystokinin: How many functions? Observations in seabreams","volume":"205","author":"Micale","year":"2014","journal-title":"Gen Comp Endocrinol"},{"key":"B47","doi-asserted-by":"publisher","first-page":"190","DOI":"10.1016\/S0167-4781(00)00263-3","article-title":"Identification and distribution of CCK-related peptides and mRNAs in the rainbow trout, Oncorhynchus mykiss1","volume":"1517","author":"Jensen","year":"2001","journal-title":"Biochim Biophys Acta BBA - Gene Struct Expr"},{"key":"B48","doi-asserted-by":"publisher","DOI":"10.1016\/0016-6480(92)90073-S","article-title":"Endocrine cells and nerves in the pyloric\u00a0ceca\u00a0and the intestine of Oncorhynchus mykiss (Teleostei): An immunocytochemical study","volume":"86","author":"Beorlegui","year":"1992","journal-title":"Gen Comp Endocrinol"},{"key":"B49","doi-asserted-by":"publisher","DOI":"10.1006\/gcen.2001.7653","article-title":"Ontogeny of cholecystokinin-immunoreactive cells in the digestive tract of atlantic halibut, hippoglossus hippoglossus, larvae","volume":"123","author":"Kamisaka","year":"2001","journal-title":"Gen Comp Endocrinol"},{"key":"B50","doi-asserted-by":"publisher","DOI":"10.1080\/00364820260294888","article-title":"Ontogeny of cholecystokinin-immunoreactive cells in the digestive tract of bluefin tuna, Thunnus thynnus, larvae","volume":"87","author":"Kamisaka","year":"2002","journal-title":"Sarsia"},{"key":"B51","doi-asserted-by":"publisher","DOI":"10.1016\/S0016-6480(03)00242-9","article-title":"Distribution of cholecystokinin-immunoreactive cells in the digestive tract of the larval teleost, ayu, Plecoglossus altivelis","volume":"134","author":"Kamisaka","year":"2003","journal-title":"Gen Comp Endocrinol"},{"key":"B52","doi-asserted-by":"publisher","DOI":"10.1016\/j.ygcen.2008.11.024","article-title":"Ghrelin, cholecystokinin, and peptide YY in Atlantic salmon (Salmo salar): Molecular cloning and tissue expression","volume":"160","author":"Murashita","year":"2009","journal-title":"Gen Comp Endocrinol"},{"key":"B53","doi-asserted-by":"publisher","DOI":"10.1016\/j.aaf.2021.11.002","article-title":"Expression of gastrin and cholecystokinin B receptor in Lateolabrax maculatus","volume":"8","author":"Cui","year":"2023","journal-title":"Aquac Fish"},{"key":"B54","doi-asserted-by":"publisher","first-page":"553","DOI":"10.3389\/fnins.2019.00553","article-title":"Effects of cholecystokinin (CCK) on gut motility in the stomachless fish ballan wrasse (Labrus bergylta)","volume":"13","author":"Le","year":"2019","journal-title":"Front Neurosci"},{"key":"B55","doi-asserted-by":"publisher","DOI":"10.2337\/diabetes.50.8.1714","article-title":"A preprandial rise in plasma ghrelin levels suggests a role in meal initiation in humans","volume":"50","author":"Cummings","year":"2001","journal-title":"Diabetes"},{"key":"B56","doi-asserted-by":"publisher","DOI":"10.1016\/j.ygcen.2009.12.006","article-title":"Plasma ghrelin concentrations change with physiological state in a sciurid hibernator (Spermophilus lateralis)","volume":"166","author":"Healy","year":"2010","journal-title":"Gen Comp Endocrinol"},{"key":"B57","doi-asserted-by":"publisher","DOI":"10.1159\/000076634","article-title":"Orexigenic actions of ghrelin in goldfish: feeding-induced changes in brain and gut mRNA expression and serum levels, and responses to central and peripheral injections","volume":"79","author":"Unniappan","year":"2004","journal-title":"Neuroendocrinology"},{"key":"B58","doi-asserted-by":"publisher","DOI":"10.1002\/jez.1784","article-title":"Molecular cloning, tissue distribution, and ontogenetic expression of ghrelin and regulation of expression by fasting and refeeding in the grass carp (Ctenopharyngodon idellus)","volume":"319","author":"Feng","year":"2013","journal-title":"J Exp Zool Part Ecol Genet Physiol"},{"key":"B59","doi-asserted-by":"publisher","DOI":"10.1016\/j.cbpa.2007.03.024","article-title":"Plasma ghrelin levels in rainbow trout in response to fasting, feeding and food composition, and effects of ghrelin on voluntary food intake","volume":"147","author":"J\u00f6nsson","year":"2007","journal-title":"Comp Biochem Physiol A Mol Integr Physiol"},{"key":"B60","doi-asserted-by":"publisher","DOI":"10.1007\/s10695-010-9434-3","article-title":"Effects of short-term starvation on ghrelin, GH-IGF system, and IGF-binding proteins in Atlantic salmon","volume":"37","author":"Hevr\u00f8y","year":"2011","journal-title":"Fish Physiol Biochem"},{"key":"B61","doi-asserted-by":"publisher","DOI":"10.1152\/ajpgi.1995.269.3.G319","article-title":"Regulation of cholecystokinin secretion by intraluminal releasing factors","volume":"269","author":"Liddle","year":"1995","journal-title":"Am J Physiol-Gastrointest Liver Physiol"},{"key":"B62","doi-asserted-by":"publisher","DOI":"10.1016\/j.ygcen.2005.09.008","article-title":"Cholecystokinin and peptide Y in yellowtail (Seriola quinqueradiata): Molecular cloning, real-time quantitative RT-PCR, and response to feeding and fasting","volume":"145","author":"Murashita","year":"2006","journal-title":"Gen Comp Endocrinol"},{"key":"B63","doi-asserted-by":"publisher","DOI":"10.1016\/j.cbpb.2006.11.009","article-title":"Changes in cholecystokinin and peptide Y gene expression with feeding in yellowtail (Seriola quinqueradiata): Relation to pancreatic exocrine regulation","volume":"146","author":"Murashita","year":"2007","journal-title":"Comp Biochem Physiol B Biochem Mol Biol"},{"key":"B64","doi-asserted-by":"publisher","first-page":"58","DOI":"10.1016\/j.yhbeh.2009.03.002","article-title":"Cloning, distribution and effects of season and nutritional status on the expression of neuropeptide Y (NPY), cocaine and amphetamine regulated transcript (CART) and cholecystokinin (CCK) in winter flounder (Pseudopleuronectes americanus)","volume":"56","author":"MacDonald","year":"2009","journal-title":"Horm Behav"},{"key":"B65","doi-asserted-by":"publisher","DOI":"10.1096\/fasebj.24.1_supplement.lb620","article-title":"Postprandial changes in GI-tract peptide hormones in the teleost Atlantic salmon","volume":"24","author":"Ronnestad","year":"2010","journal-title":"FASEB J"},{"key":"B66","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.ygcen.2013.02.003","article-title":"Molecular cloning and tissue distribution of cholecystokinin-1 receptor (CCK-1R) in yellowtail Seriola quinqueradiata and its response to feeding and in vitro CCK treatment","volume":"186","author":"Furutani","year":"2013","journal-title":"Gen Comp Endocrinol"},{"key":"B67","doi-asserted-by":"publisher","DOI":"10.1016\/j.cbpa.2010.12.006","article-title":"Cholecystokinin (CCK) in Atlantic herring (Clupea harengus L.) \u2014 Ontogeny and effects of feeding and diurnal rhythms","volume":"158","author":"Rojas-Garc\u00eda","year":"2011","journal-title":"Comp Biochem Physiol A Mol Integr Physiol"},{"key":"B68","doi-asserted-by":"publisher","DOI":"10.1016\/j.cbpa.2016.09.003","article-title":"Involvement of cholecystokinin (CCK) in the daily pattern of gastrointestinal regulation of Senegalese sole (Solea Senegalensis) larvae reared under different feeding regimes","volume":"203","author":"Navarro-Guill\u00e9n","year":"2017","journal-title":"Comp Biochem Physiol A Mol Integr Physiol"},{"key":"B69","doi-asserted-by":"publisher","DOI":"10.1016\/j.ygcen.2006.02.016","article-title":"Circulating levels of cholecystokinin and gastrin-releasing peptide in rainbow trout fed different diets","volume":"148","author":"J\u00f6nsson","year":"2006","journal-title":"Gen Comp Endocrinol"},{"key":"B70","doi-asserted-by":"publisher","DOI":"10.1152\/ajpendo.00582.2003","article-title":"Plasma ghrelin levels and hunger scores in humans initiating meals voluntarily without time- and food-related cues","volume":"287","author":"Cummings","year":"2004","journal-title":"Am J Physiol-Endocrinol Metab"},{"key":"B71","doi-asserted-by":"publisher","DOI":"10.1210\/jc.2003-031267","article-title":"Postprandial suppression of plasma ghrelin level is proportional to ingested caloric load but does not predict intermeal interval in humans","volume":"89","author":"Callahan","year":"2004","journal-title":"J Clin Endocrinol Metab"},{"key":"B72","doi-asserted-by":"publisher","DOI":"10.1016\/j.aquaculture.2025.742873","article-title":"Microalgae-derived feed additives improve physiological health, intestinal integrity, and welfare in juvenile gilthead seabream (Sparus aurata) fed plant-based diets","volume":"609","author":"Caderno","year":"2025","journal-title":"Aquaculture"},{"key":"B73","doi-asserted-by":"publisher","first-page":"76","DOI":"10.1080\/15222055.2010.549030","article-title":"Effects of partial replacement of fish meal protein by microalgae on growth, feed intake, and body composition of atlantic cod","volume":"73","author":"Walker","year":"2011","journal-title":"North Am J Aquac"},{"key":"B74","doi-asserted-by":"publisher","DOI":"10.1016\/j.heliyon.2024.e27443","article-title":"Use of grape by-products in aquaculture: New frontiers for a circular economy application","volume":"10","author":"Quagliardi","year":"2024","journal-title":"Heliyon"},{"key":"B75","doi-asserted-by":"publisher","DOI":"10.1016\/j.aqrep.2022.101248","article-title":"Biotechnological treatment of microalgae enhances growth performance, hepatic carbohydrate metabolism and intestinal physiology in gilthead seabream (Sparus aurata) juveniles close to commercial size","volume":"25","author":"Molina-Roque","year":"2022","journal-title":"Aquac Rep"},{"key":"B76","doi-asserted-by":"publisher","DOI":"10.1111\/nmo.12703","article-title":"Immunomodulatory effect of ghrelin in the intestinal mucosa","volume":"27","author":"Eissa","year":"2015","journal-title":"Neurogastroenterol Motil"},{"key":"B77","doi-asserted-by":"publisher","DOI":"10.1016\/j.cbpa.2008.05.003","article-title":"Nutrient control of release of pancreatic enzymes in yellowtail (Seriola quinqueradiata): Involvement of CCK and PY in the regulatory loop","volume":"150","author":"Murashita","year":"2008","journal-title":"Comp Biochem Physiol A Mol Integr Physiol"},{"key":"B78","doi-asserted-by":"publisher","DOI":"10.1007\/s00227-002-0790-0","article-title":"Stimulatory effect of ingested protein and\/or free amino acids on the secretion of the gastro-endocrine hormone cholecystokinin and on tryptic activity, in early-feeding herring larvae, Clupea harengus","volume":"140","author":"Koven","year":"2002","journal-title":"Mar Biol"},{"key":"B79","doi-asserted-by":"publisher","DOI":"10.1007\/s10499-008-9221-7","article-title":"Changes in the digestive enzymes and hormones of gilthead seabream larvae (Sparus aurata, L. 1758) fed on Artemia nauplii enriched with free lysine","volume":"17","author":"Naz","year":"2009","journal-title":"Aquac Int"},{"key":"B80","doi-asserted-by":"publisher","DOI":"10.1172\/JCI111809","article-title":"Cholecystokinin bioactivity in human plasma. Molecular forms, responses to feeding, and relationship to gallbladder contraction","volume":"75","author":"Liddle","year":"1985","journal-title":"J Clin Invest"},{"key":"B81","doi-asserted-by":"publisher","DOI":"10.1152\/ajpgi.1993.265.1.G107","article-title":"Evidence for indirect dietary regulation of cholecystokinin release in rats","volume":"265","author":"Sharara","year":"1993","journal-title":"Am J Physiol-Gastrointest Liver Physiol"},{"key":"B82","doi-asserted-by":"publisher","DOI":"10.1177\/153537020122601110","article-title":"Dietary protein peptic hydrolysates stimulate cholecystokinin release via direct sensing by rat intestinal mucosal cells","volume":"226","author":"Nishi","year":"2001","journal-title":"Exp Biol Med"},{"key":"B83","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1016\/j.cbpa.2016.03.010","article-title":"Daily rhythms of digestive enzyme activity and gene expression in gilthead seabream (Sparus aurata) during ontogeny","volume":"197","author":"Mata-Sotres","year":"2016","journal-title":"Comp Biochem Physiol A Mol Integr Physiol"},{"key":"B84","doi-asserted-by":"publisher","DOI":"10.1016\/j.physbeh.2004.04.013","article-title":"Endocrine controls of eating: CCK, leptin, and ghrelin","volume":"81","author":"Geary","year":"2004","journal-title":"Physiol Behav"},{"key":"B85","doi-asserted-by":"publisher","first-page":"62","DOI":"10.1016\/j.yhbeh.2017.05.004","article-title":"Ghrelin suppresses cholecystokinin (CCK), peptide YY (PYY) and glucagon-like peptide-1 (GLP-1) in the intestine, and attenuates the anorectic effects of CCK, PYY and GLP-1 in goldfish (Carassius auratus)","volume":"93","author":"Blanco","year":"2017","journal-title":"Horm Behav"}],"container-title":["Frontiers in Endocrinology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fendo.2026.1734169\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,1,27]],"date-time":"2026-01-27T05:25:23Z","timestamp":1769491523000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fendo.2026.1734169\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,1,27]]},"references-count":85,"alternative-id":["10.3389\/fendo.2026.1734169"],"URL":"https:\/\/doi.org\/10.3389\/fendo.2026.1734169","relation":{},"ISSN":["1664-2392"],"issn-type":[{"value":"1664-2392","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,1,27]]},"article-number":"1734169"}}