Gastrin

GAST
Identifikatori
Aliasi	GAST
Vanjski ID-jevi	OMIM: 137250 MGI: 104768 HomoloGene: 628 GeneCards: GAST
Lokacija gena (čovjek)
Hrom.	Hromosom 17 (čovjek)
Kraj	41,715,969 bp
Lokacija gena (miš)
Hrom.	Hromosom 11 (miš)
Kraj	100,227,822 bp
Ontologija gena
Molekularna funkcija	• GO:0001948, GO:0016582 vezivanje za proteine; • hormone activity;
Ćelijska komponenta	• extracellular region; • Vanćelijsko;
Biološki proces	• response to food; • GO:0072468 Transdukcija signala; • regulation of signaling receptor activity; • G protein-coupled receptor signaling pathway;
	Izvori:Amigo / QuickGO
Ortolozi
	2520
	14459
	ENSG00000184502
	ENSMUSG00000017165
	P01350
	P48757
	NM_000805
	NM_010257
	NP_000796
	NP_034387
	Wikipodaci
Pogledaj/uredi – čovjek	Pogledaj/uredi – miš

Gastrin je peptidni hormon koji stimulira lučenje želučane kiseline (HCl) u parijetalnim ćelijama želuca i pomaže u njegovoj pokretljivosti. Oslobađaju ga G-ćelije u pilornom antrumu želuca, dvanaestopalačnom crijevu i gušterači.

Gastrin se veže za holecistokininski B receptor da stimulira oslobađanje histamina u enterohromafinu-sličnim ćelijama i inducira umetanje K⁺/ H⁺ ATPaza pumpi u apikalnu membranu parijetalnih ćelija (što zauzvrat povećava otpuštanje H⁺ u želudačnu šupljinu). Njegovo oslobađanje stimuliraju peptidi u lumenu želuca. U ljudi, gen GAS nalazi se na dugom kraku hromosoma 17, pozicija 17q21).^[5]

Fiziologija

Snteza

Gastrin je linearni peptidni hormon koji proizvode G-ćelije dvanaestopalačnog crijeva i pilorni antrum želuca. Izlučuje se u krvotok. Kodirani polipeptid je preprogastrin, koji se enzimima cijepa u posttranslacijskim modifikacijama dajući progastrin (međuprodukt, neaktivni prekursor), a zatim gastrin u raznim oblicima, prvenstveno sljedeća tri:

Također, pentagastrin je veštački sintetizirana petoaminokiselinska sekvenca, sa identičnim posljednjih pet aminokiselinskih sekvenci na C-kraju gastrina. Brojevi se odnose na broj aminokiselina.

Lučenje

Gastrin se oslobađa kao odgovor na određene podražaje. Oni uključuju:

antrum želuca distenzija
Vagus nustimulaciju (posredovana signalna ćelija neurokrinim bombezinom ili GRP-om kod ljudi)
prisustvo delimične digestije proteina, posebno aminokiselina, u želucu.

Aromatske aminokiseline su posebno snažni podražaji za oslobađanje gastrina;^[6]

hiperkalcemija (preko kalcij-osjetljivih receptora.^[7])

Lučenje gastrina inhibiraju:^[8]^[9]

prisustvo kiselina (prvenstveno lučene HCl) u želucu (slučaj negativne povratne informacije)
somatostatin takođe inhibira oslobađanje gastrina, zajedno sa sekretinom, GIP (gastroinhibitorni peptid), VIP (vazoaktivni crijevni peptid), glukagon i kalcitonin.

Funkcija

G-ćelija vidljiva je u donjem lijevom uglu, a gastrin je označen kao dve crne strelice koje vode od njega. Napomena: ovaj dijagram ne ilustrira stimulativni efekat gastrina na ECL ćelije.

Prisustvo gastrina stimulira parijetalne ćelije želuca da luče hlorovodoničnu kiselinu (HCl) / želučanu kiselinu. To se odvija direktno. u parijetalnim ćelijama i indirektno, vezanjem na CCK2 / gastrinski receptori na ECL-ćelijama u želucu, koji zatim reagira oslobađanjem histamina, koji zauzvrat djeluje parakrino na parijetalne ćelije, stimulirajući ih da luče H + ion. Ovo je glavni podsticaj za sekreciju kiseline u parijetalnim ćelijama.^[10]

Zajedno sa gore spomenutom funkcijom, pokazalo se da gastrin ima i dodatne aktivnosti:

Podstiče sazrijevanje parijetalnih ćelija i rast fundusa.
Uzrokuje da glavne ćelije da luče pepsinogen, zimogen (neaktivan) oblik probavnog enzima pepsina.
Povećava pokretljivost antralnih mišića i pospješuje kontrakcije želuca.
Jača antralne kontrakcije protiv pilorusa i opušta pilorični sfinkter, što povećava brzinu pražnjenja želuca.^[11]
Plays a role in the relaxation of the ileocecal valve.^[12]
Izaziva sekreciju pankreasa i pražnjenje žučne kese.^[13]
Može uticati na tonus donji jednjački sfinkter (LES), uzrokujući njegovu kontrakciju,^[14] iako uzrok može biti i pentagastrin, a ne endogeni gastrin.^[15]
Gastrin doprinosi pojavi gastrokoličnog refleksa.

Faktori koji utiču na sekreciju

Faktori koji utiču na lučenje gastrina mogu se podijeliti u dvije kategorije:^[16]

Fiziološki

Lumen želuca

Stimulativni faktori: hranjivi proteini i aminokiseline (meso), hiperkalcemija. (npr. tokom želučane faze)
Inhibitorni faktori: kiselost (pH ispod 3) – mehanizam negativne povratne sprege, oslobađanjem somatostatina iz δ ćelija u želucu, koji inhibira oslobađanje gastrina i histamina.

Parakrini

Stimulativni faktori: bombesin ili gastrin-oslobađajući peptid (GRP)
Inhibitorni faktori: somatostatin – djeluje parakrino na somatostatin-2 receptore na G-ćelijama, putem lokalne difuzije u međućelijskim prostorima, ali i sistemski kroz njegovo puštanje u lokalnu cirkulaciju sluznice; inhibira lučenje kiseline djelujući na parijetalne ćelije.

Nervni

Stimulativni faktori: Beta-adrenergični agensi, holinergijski agensi, gastrin-oslobađajući peptid (GRP)
Inhibitorni faktor: Enterogastrični refleks

Cirkulacijski

Stimulacijski faktor: Gastrin;
Inhibitorni faktori: Želučani inhibitorni peptid (GIP), sekretin, somatostatin, glukagon, kalcitonin.

Patofiziološki

Paraneoplazijski

Prekomjerna sekrecija gastrina izaziva paraneoplazijski sindrom.

Uloga u bolesti

Kod osoba sa Zollinger-Ellisonovim sindromom, gastrin se proizvodi na prekomjernoj razini, često preko gastrinoma (tumor koji proizvodi gastrin, uglavnom benigni) dvanaestpalačnog crijeva ili pankreasa.^[17]

U autoimunskom gastritisu, imunoski sistem napada parijetalne ćelije, što dovodi do hipohlorhidrije (niskog lučenja želučane kiseline). To rezultira povišenom razinom gastrina, u pokušaju da se podesi povećani pH u želucu. Na kraju se sve parijetalne ćelije izgube, pa aklorhidrija rezultira gubitkom negativne povratne informacije na sekreciju gastrina. Koncentracija gastrina u plazmi povišena je kod gotovo svih osoba sa poremećajem zvanim mukolipidoza tip IV (srednja vrednost 1507 pg/ml; raspon 400-4100 pg/ml) (normala 0-200 pg /ml), sekundarno u odnosu na konstitutivnu aklorhidriju. Ovo otkriće olakšava dijagnozu kod pacijenata sa ovim neurogenetičkim poremećajemIn autoimunskim gastritisom.^[18] Pored toga, povišeni nivoi gastrina mogu biti prisutni pri hroničnom gastritisu koji je posljedica infekcije sa H. pylori.^[19]

Historija

Postojanje gastrina prvi je put predložio britanski fiziolog John Sydney Edkins, 1905.,^[20]^[21] a gastrine su izolirali Hilda Tracy i Roderic Alfred Gregory sa University of Liverpool, 1964. godine.^[22] In 1964 the structure of gastrin was determined.^[23]

Reference

^ ^a ^b ^c GRCh38: Ensembl release 89: ENSG00000184502 - Ensembl, maj 2017
^ ^a ^b ^c GRCm38: Ensembl release 89: ENSMUSG00000017165 - Ensembl, maj 2017
^ "Human PubMed Reference:". National Center for Biotechnology Information, U.S. National Library of Medicine.
^ "Mouse PubMed Reference:". National Center for Biotechnology Information, U.S. National Library of Medicine.
^ Lund T, Geurts van Kessel AH, Haun S, Dixon JE (maj 1986). "The genes for human gastrin and cholecystokinin are located on different chromosomes". Human Genetics. 73 (1): 77–80. doi:10.1007/BF00292669. PMID 3011648.
^ Blanco, Antonio; Blanco, Gustavo (2017), "Biochemical Bases of Endocrinology (II) Hormones and Other Chemical Intermediates", Medical Biochemistry, Elsevier, str. 573–644, doi:10.1016/b978-0-12-803550-4.00026-4, ISBN 9780128035504
^ Feng J, Petersen CD, Coy DH, Jiang JK, Thomas CJ, Pollak MR, Wank SA (oktobar 2010). "Calcium-sensing receptor is a physiologic multimodal chemosensor regulating gastric G-cell growth and gastrin secretion". Proceedings of the National Academy of Sciences of the United States of America. 107 (41): 17791–6. doi:10.1073/pnas.1009078107. PMC 2955134. PMID 20876097.
^ Holst JJ, Orskov C, Seier-Poulsen S (1992). "Somatostatin is an essential paracrine link in acid inhibition of gastrin secretion". Digestion. 51 (2): 95–102. doi:10.1159/000200882. PMID 1354190.
^ Johnson LR (mart 1984). "Effects of somatostatin and acid on inhibition of gastrin release in newborn rats". Endocrinology. 114 (3): 743–6. doi:10.1210/endo-114-3-743. PMID 6141932. Arhivirano s originala, 2008-09-0. Pristupljeno 17. 5. 2011. Provjerite vrijednost datuma u parametru: |archive-date= (pomoć)
^ Lindström, E.; Chen, D.; Norlén, P.; Andersson, K.; Håkanson, R. (2001). "Control of gastric acid secretion:the gastrin-ECL cell-parietal cell axis". Comparative Biochemistry and Physiology. Part A, Molecular & Integrative Physiology. 128 (3): 505–514. doi:10.1016/s1095-6433(00)00331-7. ISSN 1095-6433. PMID 11246041.
^ Tortora, G. J., & Grabowski, S. R. (1996). Principles of anatomy and physiology. New York, NY: HarperCollins College. 14th Ed. Pg 906
^ Vadokas B, Lüdtke FE, Lepsien G, Golenhofen K, Mandrek K (decembar 1997). "Effects of gastrin-releasing peptide (GRP) on the mechanical activity of the human ileocaecal region in vitro". Neurogastroenterology and Motility. 9 (4): 265–70. doi:10.1046/j.1365-2982.1997.d01-59.x. PMID 9430795.
^ Valenzuela JE, Walsh JH, Isenberg JI (septembar 1976). "Effect of gastrin on pancreatic enzyme secretion and gallbladder emptying in man". Gastroenterology. 71 (3): 409–11. doi:10.1016/S0016-5085(76)80445-3. PMID 950091.
^ Castell DO (februar 1978). "Gastrin and lower esophageal sphincter tone". Archives of Internal Medicine. 138 (2): 196. doi:10.1001/archinte.138.2.196. PMID 626547.
^ Henderson JM, Lidgard G, Osborne DH, Carter DC, Heading RC (februar 1978). "Lower oesophageal sphincter response to gastrin--pharmacological or physiological?". Gut. 19 (2): 99–102. doi:10.1136/gut.19.2.99. PMC 1411818. PMID 631634.
^ Indu Khurana (2006). Textbook medical physiology. New Delhi: Reed Elsevier India. str. 605. ISBN 978-8181478504. OCLC 968478170.
^ Baron, J. H. (1978). Clinical Tests of Gastric Secretion (jezik: engleski). doi:10.1007/978-1-349-03188-7. ISBN 978-1-349-03190-0.
^ Schiffmann R, Dwyer NK, Lubensky IA, Tsokos M, Sutliff VE, Latimer JS, Frei KP, Brady RO, Barton NW, Blanchette-Mackie EJ, Goldin E (februar 1998). "Constitutive achlorhydria in mucolipidosis type IV". Proceedings of the National Academy of Sciences of the United States of America. 95 (3): 1207–12. doi:10.1073/pnas.95.3.1207. PMC 18720. PMID 9448310.
^ "Review Article: Strategies to Determine Whether Hypergastrinaemia Is Due to Zollinger-Ellison Syndrome Rather Than a More Common Benign Cause". www.medscape.com. Arhivirano s originala, 22. 5. 2023. Pristupljeno 2. 9. 2020.
^ Edkins JS (mart 1906). "The chemical mechanism of gastric secretion". The Journal of Physiology. 34 (1–2): 133–44. doi:10.1113/jphysiol.1906.sp001146. PMC 1465807. PMID 16992839. Arhivirano s originala, 7. 2. 2020. Pristupljeno 2. 9. 2020.
^ Modlin IM, Kidd M, Marks IN, Tang LH (februar 1997). "The pivotal role of John S. Edkins in the discovery of gastrin". World Journal of Surgery. 21 (2): 226–34. doi:10.1007/s002689900221. PMID 8995084.
^ Gregory RA, Tracy HJ (1964). "The constitution and properties of two gastrins extracted from hog antral mucosa: Part I the isolation of two gastrins from hog antral mucosa". Gut. 5 (2): 103–107. doi:10.1136/gut.5.2.103. PMC 1552180. PMID 14159395. CS1 održavanje: nepreporučeni parametar (link)
^ Gregory H, Hardy PM, Jones DS, Kenner GW, Sheppard RC (decembar 1964). "The antral hormone gastrin. Structure of gastrin". Nature. 204 (4962): 931–3. doi:10.1038/204931a0. PMID 14248711.

Dopunska literatura

Rozengurt E, Walsh JH (2001). "Gastrin, CCK, signaling, and cancer". Annual Review of Physiology. 63: 49–76. doi:10.1146/annurev.physiol.63.1.49. PMID 11181948.
Dockray GJ (decembar 2004). "Clinical endocrinology and metabolism. Gastrin". Best Practice & Research. Clinical Endocrinology & Metabolism. 18 (4): 555–68. doi:10.1016/j.beem.2004.07.003. PMID 15533775.
Anlauf M, Garbrecht N, Henopp T, Schmitt A, Schlenger R, Raffel A, Krausch M, Gimm O, Eisenberger CF, Knoefel WT, Dralle H, Komminoth P, Heitz PU, Perren A, Klöppel G (septembar 2006). "Sporadic versus hereditary gastrinomas of the duodenum and pancreas: distinct clinico-pathological and epidemiological features". World Journal of Gastroenterology. 12 (34): 5440–6. doi:10.3748/wjg.v12.i34.5440. PMC 4088224. PMID 17006979.
Polosatov MV, Klimov PK, Masevich CG, Samartsev MA, Wünsch E (april 1979). "Interaction of synthetic human big gastrin with blood proteins of man and animals". Acta Hepato-Gastroenterologica. 26 (2): 154–9. PMID 463490.
Fritsch WP, Hausamen TU, Scholten T (april 1977). "[Gastrointestinal hormones. I. Hormones of the gastrin group]". Zeitschrift für Gastroenterologie. 15 (4): 264–76. PMID 871064.
Higashimoto Y, Himeno S, Shinomura Y, Nagao K, Tamura T, Tarui S (maj 1989). "Purification and structural determination of urinary NH2-terminal big gastrin fragments". Biochemical and Biophysical Research Communications. 160 (3): 1364–70. doi:10.1016/S0006-291X(89)80154-8. PMID 2730647.
Pauwels S, Najdovski T, Dimaline R, Lee CM, Deschodt-Lanckman M (juni 1989). "Degradation of human gastrin and CCK by endopeptidase 24.11: differential behaviour of the sulphated and unsulphated peptides". Biochimica et Biophysica Acta (BBA) - Protein Structure and Molecular Enzymology. 996 (1–2): 82–8. doi:10.1016/0167-4838(89)90098-8. PMID 2736261.
Lund T, Geurts van Kessel AH, Haun S, Dixon JE (maj 1986). "The genes for human gastrin and cholecystokinin are located on different chromosomes". Human Genetics. 73 (1): 77–80. doi:10.1007/BF00292669. PMID 3011648.
Kariya Y, Kato K, Hayashizaki Y, Himeno S, Tarui S, Matsubara K (1986). "Expression of human gastrin gene in normal and gastrinoma tissues". Gene. 50 (1–3): 345–52. doi:10.1016/0378-1119(86)90338-0. PMID 3034736.
Gregory RA, Tracy HJ, Agarwal KL, Grossman MI (august 1969). "Aminoacid constitution of two gastrins isolated from Zollinger-Ellison tumour tissue". Gut. 10 (8): 603–8. doi:10.1136/gut.10.8.603. PMC 1552899. PMID 5822140.
Bentley PH, Kenner GW, Sheppard RC (februar 1966). "Structures of human gastrins I and II". Nature. 209 (5023): 583–5. doi:10.1038/209583b0. PMID 5921183.
Ito R, Sato K, Helmer T, Jay G, Agarwal K (august 1984). "Structural analysis of the gene encoding human gastrin: the large intron contains an Alu sequence". Proceedings of the National Academy of Sciences of the United States of America. 81 (15): 4662–6. doi:10.1073/pnas.81.15.4662. PMC 391550. PMID 6087340.
Wiborg O, Berglund L, Boel E, Norris F, Norris K, Rehfeld JF, Marcker KA, Vuust J (februar 1984). "Structure of a human gastrin gene". Proceedings of the National Academy of Sciences of the United States of America. 81 (4): 1067–9. doi:10.1073/pnas.81.4.1067. PMC 344765. PMID 6322186.
Kato K, Hayashizaki Y, Takahashi Y, Himeno S, Matsubara K (decembar 1983). "Molecular cloning of the human gastrin gene". Nucleic Acids Research. 11 (23): 8197–203. doi:10.1093/nar/11.23.8197. PMC 326575. PMID 6324077.
Boel E, Vuust J, Norris F, Norris K, Wind A, Rehfeld JF, Marcker KA (maj 1983). "Molecular cloning of human gastrin cDNA: evidence for evolution of gastrin by gene duplication". Proceedings of the National Academy of Sciences of the United States of America. 80 (10): 2866–9. doi:10.1073/pnas.80.10.2866. PMC 393933. PMID 6574456.
Kato K, Himeno S, Takahashi Y, Wakabayashi T, Tarui S, Matsubara K (decembar 1983). "Molecular cloning of human gastrin precursor cDNA". Gene. 26 (1): 53–7. doi:10.1016/0378-1119(83)90035-5. PMID 6689486.
Koh TJ, Wang TC (novembar 1995). "Molecular cloning and sequencing of the murine gastrin gene". Biochemical and Biophysical Research Communications. 216 (1): 34–41. doi:10.1006/bbrc.1995.2588. PMID 7488110.
Rehfeld JF, Hansen CP, Johnsen AH (januar 1995). "Post-poly(Glu) cleavage and degradation modified by O-sulfated tyrosine: a novel post-translational processing mechanism". The EMBO Journal. 14 (2): 389–96. doi:10.1002/j.1460-2075.1995.tb07013.x. PMC 398093. PMID 7530658.
Rehfeld JF, Johnsen AH (august 1994). "Identification of gastrin component I as gastrin-71. The largest possible bioactive progastrin product". European Journal of Biochemistry. 223 (3): 765–73. doi:10.1111/j.1432-1033.1994.tb19051.x. PMID 8055952.
Varro A, Dockray GJ (novembar 1993). "Post-translational processing of progastrin: inhibition of cleavage, phosphorylation and sulphation by brefeldin A". The Biochemical Journal. 295 (Pt 3): 813–9. doi:10.1042/bj2950813. PMC 1134634. PMID 8240296.

Vanjski linkovi

Overview at colostate.edu Arhivirano 19. 5. 2017. na Wayback Machine
Nosek, Thomas M. "Section 6/6ch4/s6ch4_14". Essentials of Human Physiology. Arhivirano s originala, 24. 3. 2016.

Commons logo

Commons ima datoteke na temu: Gastrin

[refGRCh38Ensembl-1] GRCh38: Ensembl release 89: ENSG00000184502 - Ensembl, maj 2017

[refGRCm38Ensembl-2] GRCm38: Ensembl release 89: ENSMUSG00000017165 - Ensembl, maj 2017

[3] "Human PubMed Reference:". National Center for Biotechnology Information, U.S. National Library of Medicine.

[4] "Mouse PubMed Reference:". National Center for Biotechnology Information, U.S. National Library of Medicine.

[5] Lund T, Geurts van Kessel AH, Haun S, Dixon JE (maj 1986). "The genes for human gastrin and cholecystokinin are located on different chromosomes". Human Genetics. 73 (1): 77–80. doi:10.1007/BF00292669. PMID 3011648.

[6] Blanco, Antonio; Blanco, Gustavo (2017), "Biochemical Bases of Endocrinology (II) Hormones and Other Chemical Intermediates", Medical Biochemistry, Elsevier, str. 573–644, doi:10.1016/b978-0-12-803550-4.00026-4, ISBN 9780128035504

[7] Feng J, Petersen CD, Coy DH, Jiang JK, Thomas CJ, Pollak MR, Wank SA (oktobar 2010). "Calcium-sensing receptor is a physiologic multimodal chemosensor regulating gastric G-cell growth and gastrin secretion". Proceedings of the National Academy of Sciences of the United States of America. 107 (41): 17791–6. doi:10.1073/pnas.1009078107. PMC 2955134. PMID 20876097.

[8] Holst JJ, Orskov C, Seier-Poulsen S (1992). "Somatostatin is an essential paracrine link in acid inhibition of gastrin secretion". Digestion. 51 (2): 95–102. doi:10.1159/000200882. PMID 1354190.

[9] Johnson LR (mart 1984). "Effects of somatostatin and acid on inhibition of gastrin release in newborn rats". Endocrinology. 114 (3): 743–6. doi:10.1210/endo-114-3-743. PMID 6141932. Arhivirano s originala, 2008-09-0. Pristupljeno 17. 5. 2011. Provjerite vrijednost datuma u parametru: |archive-date= (pomoć)

[10] Lindström, E.; Chen, D.; Norlén, P.; Andersson, K.; Håkanson, R. (2001). "Control of gastric acid secretion:the gastrin-ECL cell-parietal cell axis". Comparative Biochemistry and Physiology. Part A, Molecular & Integrative Physiology. 128 (3): 505–514. doi:10.1016/s1095-6433(00)00331-7. ISSN 1095-6433. PMID 11246041.

[11] Tortora, G. J., & Grabowski, S. R. (1996). Principles of anatomy and physiology. New York, NY: HarperCollins College. 14th Ed. Pg 906

[pmid9430795-12] Vadokas B, Lüdtke FE, Lepsien G, Golenhofen K, Mandrek K (decembar 1997). "Effects of gastrin-releasing peptide (GRP) on the mechanical activity of the human ileocaecal region in vitro". Neurogastroenterology and Motility. 9 (4): 265–70. doi:10.1046/j.1365-2982.1997.d01-59.x. PMID 9430795.

[pmid950091-13] Valenzuela JE, Walsh JH, Isenberg JI (septembar 1976). "Effect of gastrin on pancreatic enzyme secretion and gallbladder emptying in man". Gastroenterology. 71 (3): 409–11. doi:10.1016/S0016-5085(76)80445-3. PMID 950091.

[pmid626547-14] Castell DO (februar 1978). "Gastrin and lower esophageal sphincter tone". Archives of Internal Medicine. 138 (2): 196. doi:10.1001/archinte.138.2.196. PMID 626547.

[pmid631634-15] Henderson JM, Lidgard G, Osborne DH, Carter DC, Heading RC (februar 1978). "Lower oesophageal sphincter response to gastrin--pharmacological or physiological?". Gut. 19 (2): 99–102. doi:10.1136/gut.19.2.99. PMC 1411818. PMID 631634.

[16] Indu Khurana (2006). Textbook medical physiology. New Delhi: Reed Elsevier India. str. 605. ISBN 978-8181478504. OCLC 968478170.

[17] Baron, J. H. (1978). Clinical Tests of Gastric Secretion (jezik: engleski). doi:10.1007/978-1-349-03188-7. ISBN 978-1-349-03190-0.

[pmid9448310-18] Schiffmann R, Dwyer NK, Lubensky IA, Tsokos M, Sutliff VE, Latimer JS, Frei KP, Brady RO, Barton NW, Blanchette-Mackie EJ, Goldin E (februar 1998). "Constitutive achlorhydria in mucolipidosis type IV". Proceedings of the National Academy of Sciences of the United States of America. 95 (3): 1207–12. doi:10.1073/pnas.95.3.1207. PMC 18720. PMID 9448310.

[19] "Review Article: Strategies to Determine Whether Hypergastrinaemia Is Due to Zollinger-Ellison Syndrome Rather Than a More Common Benign Cause". www.medscape.com. Arhivirano s originala, 22. 5. 2023. Pristupljeno 2. 9. 2020.

[20] Edkins JS (mart 1906). "The chemical mechanism of gastric secretion". The Journal of Physiology. 34 (1–2): 133–44. doi:10.1113/jphysiol.1906.sp001146. PMC 1465807. PMID 16992839. Arhivirano s originala, 7. 2. 2020. Pristupljeno 2. 9. 2020.

[21] Modlin IM, Kidd M, Marks IN, Tang LH (februar 1997). "The pivotal role of John S. Edkins in the discovery of gastrin". World Journal of Surgery. 21 (2): 226–34. doi:10.1007/s002689900221. PMID 8995084.

[22] Gregory RA, Tracy HJ (1964). "The constitution and properties of two gastrins extracted from hog antral mucosa: Part I the isolation of two gastrins from hog antral mucosa". Gut. 5 (2): 103–107. doi:10.1136/gut.5.2.103. PMC 1552180. PMID 14159395. CS1 održavanje: nepreporučeni parametar (link)

[pmid14248711-23] Gregory H, Hardy PM, Jones DS, Kenner GW, Sheppard RC (decembar 1964). "The antral hormone gastrin. Structure of gastrin". Nature. 204 (4962): 931–3. doi:10.1038/204931a0. PMID 14248711.

[1]