胃促生長素曹曙光，鄭君傑，薛戰雄。胃促生長素與消化道運動 J 國際消化病雜誌，200727 4290292（，lenomorelin）全名胃生長激素釋放肽（growth hormonereleasing peptide）王馨苒，賈許楊，杜磊，等。胃生長激素釋放肽調控食慾的機制及臨床應用前景 J 中華內分泌代謝雜誌，2023 39 3 283288 DOI 103760cmajcn3112822022070900421，俗稱飢餓素（hunger hormone），是一種主要由胃腸道（尤其是胃）的產生的激素，屬一種多功能腦腸肽（胃腸道神經肽）Kojima M Hosoda H Date Y Nakazato M Matsuo H Kangawa K December 1999 Ghrelin is a growthhormonereleasing acylated peptide from stomach Nature 402 6762 65660 Bibcode1999Natur402656K doi10103845230 PMID 10604470Müller TD Nogueiras R Andermann ML Andrews ZB Anker SD Argente J et al June 2015 Ghrelin Molecular Metabolism 4 6 43760 doi101016jmolmet201503005 PMC 4443295 PMID 26042199，能誘導生長激素的釋放、胃腸道功能調節、攝食、抑制炎症反應，並維持能量正平衡的作用。 它被稱為飢餓素是因為它會增加進食的動力，在餐前飢餓時，飢餓素在血液中濃度最高，進食後恢復到較低水平 Cummings DE Purnell JQ Frayo RS Schmidova K Wisse BE Weigle DS August 2001 A preprandial rise in plasma ghrelin levels suggests a role in meal initiation in humans Diabetes 50 8 17149 doi102337diabetes5081714 PMID 11473029。飢餓素可能透過增加胃的活動性和刺激胃酸分泌來幫助進食準備 Schwartz MW Woods SC Porte D Seeley RJ Baskin DG April 2000 Central nervous system control of food intake Nature 404 6778 66171 doi10103835007534 PMID 10766253。 飢餓素能夠活化腦垂體前葉和下視丘弓狀核中的細胞，Dickson SL Leng G Robinson IC March 1993 Systemic administration of growth hormonereleasing peptide activates hypothalamic arcuate neurons Neuroscience 53 2 3036 doi10101603064522 93 90197n PMID 8492908 其中包括可引起食慾的神經肽 Y 神經元。Dickson SL Luckman SM February 1997 Induction of cfos messenger ribonucleic acid in neuropeptide Y and growth hormone GHreleasing factor neurons in the rat arcuate nucleus following systemic injection of the GH secretagogue GHreleasing peptide6 Endocrinology 138 2 7717 doi101210endo13824907 PMID 9003014 飢餓素能夠刺激具有特定受體的大腦結構：飢餓素受體 1A（GHSR 1A）。Howard AD Feighner SD Cully DF Arena JP Liberator PA Rosenblum CI et al August 1996 A receptor in pituitary and hypothalamus that functions in growth hormone release Science 273 5277 9747 Bibcode1996Sci273974H doi101126science2735277974 PMID 8688086 飢餓素同時也能夠調節神經結構中的犒賞系統 Nestler EJ Hyman SE Holtzman DM Malenka RC 2015 Neural and Neuroendocrine Control of the Internal Milieu Molecular Neuropharmacology A Foundation for Clinical Neuroscience 3rd ed New York McGrawHill Medical pp 245267 ISBN 9780071827690、學習和記憶，睡眠 甦醒週期、味覺、犒賞行為和葡萄糖代謝。Dickson SL Egecioglu E Landgren S Skibicka KP Engel JA Jerlhag E June 2011 The role of the central ghrelin system in reward from food and chemical drugs Molecular and Cellular Endocrinology 340 1 807 doi101016jmce201102017 hdl207726318 PMID 21354264Le Moal M 2002 Mesocorticolimbic Dopaminergic Neurons In Davis KL Charney D Coyle JT Nemeroff C eds Neuropsychopharmacology the fifth generation of progress an official publication of the American College of Neuropsychopharmacology 5th ed Philadelphia Pa Lippincott Williams Wilkins ISBN 9780781728379 歷史與命名 飢餓素的首次發現是在 1999 年發現了飢餓素受體（稱為生長激素促分泌素受體 1A 或 GHSR）之後。其命名是以它能夠「促生長激素分泌」的功能來命名，源自於原始印歐語中的 gʰre，意謂成長。 基因、轉錄產物與構造 其 GHRL 基因產生的 mRNA 有四個外顯子，並產生五種產物：第一種是 117 個氨基酸的前飢餓素原。（與促胃動素原同源；兩者都是促胃動素家族的成員）。將前飢餓素原剪接後產生飢餓素原，再剪接後產生一個具有 28 個氨基酸的未醯化飢餓素和一個醯化飢餓素（CGherlin）。據推測，肥胖抑制素可能從 Cghrelin 剪接而成。Seim I Amorim L Walpole C Carter S Chopin LK Herington AC January 2010 Ghrelin generelated peptides multifunctional endocrine autocrine modulators in health and disease Clinical and Experimental Pharmacology Physiology 37 1 12531 doi101111j14401681200905241x PMID 19566830 飢餓素只有在飢餓素醯基轉移酶 （GOAT）的幫助下，結構中的辛酸在轉譯後與絲氨酸的 3 號位連線時才具有活性。它位於胃和胰臟中的飢餓素細胞的細胞膜上 Castañeda TR Tong J Datta R Culler M Tschöp MH January 2010 Ghrelin in the regulation of body weight and metabolism Frontiers in Neuroendocrinology 31 1 4460 doi101016jyfrne200910008 PMID 19896496，而未辛醯化的形式則稱為去醯基飢餓素，其無法活化 GHSR 受體，但具有其他作用：心臟方面的功能 Bedendi I Alloatti G Marcantoni A Malan D Catapano F Ghé C et al August 2003 Cardiac effects of ghrelin and its endogenous derivatives desoctanoyl ghrelin and desGln14ghrelin PDF European Journal of Pharmacology 476 12 8795 doi101016S00142999 03 020831 hdl2318125949 PMID 12969753、刺激食慾 Toshinai K Yamaguchi H Sun Y Smith RG Yamanaka A Sakurai T et al May 2006 Desacyl ghrelin induces food intake by a mechanism independent of the growth hormone secretagogue receptor Endocrinology 147 5 230614 doi101210en20051357 PMID 16484324 和抑制肝臟輸出葡萄糖 Gauna C Delhanty PJ Hofland LJ Janssen JA Broglio F Ross RJ et al February 2005 Ghrelin stimulates whereas desoctanoyl ghrelin inhibits glucose output by primary hepatocytes The Journal of Clinical Endocrinology and Metabolism 90 2 105560 doi101210jc20041069 PMID 15536157。同時也觀察到，辛醯基以外的側鏈也可以觸發飢餓素受體 Korbonits M Goldstone AP Gueorguiev M Grossman AB April 2004 Ghrelina hormone with multiple functions Frontiers in Neuroendocrinology 25 1 2768 doi101016jyfrne200403002 PMID 15183037。 特別的是，已發現癸醯飢餓素在小鼠迴圈中的飢餓素站了一定比例，但截至 2011 年，其在人體中的存在尚未確定。Stengel A Taché Y June 2011 Interaction between gastric and upper small intestinal hormones in the regulation of hunger and satiety ghrelin and cholecystokinin take the central stage Current Protein Peptide Science 12 4 293304 doi102174138920311795906673 PMC 3670092 PMID 21428875 飢餓素細胞 別名 飢餓素細胞也被稱為胰臟的 Alike cell、Xcell（X 意謂功能不明）、小鼠中的 XAlike cell、胰臟中的 ε 細胞、人體的 PD sub 1 cell、或 Gr cell（Gherlin 的簡稱）Zigman JM Nakano Y Coppari R Balthasar N Marcus JN Lee CE et al December 2005 Mice lacking ghrelin receptors resist the development of dietinduced obesity The Journal of Clinical Investigation 115 12 356472 doi101172JCI26002 PMC 1297251 PMID 16322794。 位置 飢餓素細胞主要存在於胃 Ariyasu H Takaya K Tagami T Ogawa Y Hosoda K Akamizu T et al October 2001 Stomach is a major source of circulating ghrelin and feeding state determines plasma ghrelinlike immunoreactivity levels in humans The Journal of Clinical Endocrinology and Metabolism 86 10 47538 doi101210jcem86107885 PMID 11600536 和十二指腸中，但也存在於空腸、肺、胰島 Suckale J Solimena M May 2008 Pancreas islets in metabolic signalingfocus on the betacell Frontiers in Bioscience 13 13 715671 doi1027413218 PMID 18508724、性腺、腎上腺皮質、胎盤和腎臟中。最近也有研究顯示，飢餓素能夠在大腦區域性產生 Ferrini F Salio C Lossi L Merighi A March 2009 Ghrelin in central neurons Current Neuropharmacology 7 1 3749 doi102174157015909787602779 PMC 2724662 PMID 19721816。 特色 飢餓素細胞存在於胃底腺細胞中（佔細胞的 20％）Simonsson M Eriksson S Håkanson R Lind T Lönroth H Lundell L et al November 1988 Endocrine cells in the human oxyntic mucosa A histochemical study Scandinavian Journal of Gastroenterology 23 9 108999 doi10310900365528809090174 PMID 2470131、幽門和小腸中，是卵圓形的顆粒性細胞 Grube D Forssmann WG November 1979 Morphology and function of the enteroendocrine cells Hormone and Metabolic Research Hormon und Stoffwechselforschung Hormones et Metabolisme 11 11 589606 doi101055s00281092785 PMID 94030，同時具有胃泌素受體 Fukumoto K Nakahara K Katayama T Miyazatao M Kangawa K Murakami N September 2008 Synergistic action of gastrin and ghrelin on gastric acid secretion in rats Biochemical and Biophysical Research Communications 374 1 603 doi101016jbbrc200806114 PMID 18611393，且有一些能夠產生脂肪激素 nesfatin1Inhoff T Stengel A Peter L Goebel M Taché Y Bannert N et al February 2010 Novel insight in distribution of nesfatin1 and phosphomTOR in the arcuate nucleus of the hypothalamus of rats Peptides 31 2 25762 doi101016jpeptides200911024 PMC 4043136 PMID 19961888。飢餓素細胞在胰臟中沒有終末分化的步驟，它們作為前驅細胞，可以產生 A 細胞、PP 細胞和 β 細胞。Arnes L Hill JT Gross S Magnuson MA Sussel L 2012 Ghrelin expression in the mouse pancreas defines a unique multipotent progenitor population PLOS ONE 7 12 e52026 Bibcode2012PLoSO752026A doi101371journalpone0052026 PMC 3520898 PMID 23251675 作用的功能與機制 飢餓素參與調節的複雜過程，透過調節飢餓訊號來影響能量輸入；調節 ATP 生產、脂肪儲存、肝醣儲存及短期熱消耗等能量比例調節來影響能量輸出。能量平衡的結果最終反映在體重上，並根據代謝訊號和需求進行持續監測調整。而胃腦溝通是影響能量平衡的重要途徑，有幾種溝通路徑，其中包括胃細胞內 路徑所調節的飢餓素、及胃系統彼此間的互動作用，Folgueira C Seoane LM Casanueva FF 2014 The brainstomach connection In Delhanty PJD van der Lely AJ eds How Gut and Brain Control Metabolism Frontiers of Hormone Research 42 Basel Karger pp 8392 doi101159000358316 ISBN 9783318026382 PMID 24732927 及迷走神經的傳出入訊號。 飢餓素及合成飢餓素模擬物 透過誘發含及 Chen HY Trumbauer ME Chen AS Weingarth DT Adams JR Frazier EG et al June 2004 Orexigenic action of peripheral ghrelin is mediated by neuropeptide Y and agoutirelated protein Endocrinology 145 6 260712 doi101210en20031596 PMID 14962995 的弓形核受體來增加體重及脂肪量。Lall S Tung LY Ohlsson C Jansson JO Dickson SL January 2001 Growth hormone GHindependent stimulation of adiposity by GH secretagogues Biochemical and Biophysical Research Communications 280 1 1328 doi101006bbrc20004065 PMID 11162489Tschöp M Smiley DL Heiman ML October 2000 Ghrelin induces adiposity in rodents Nature 407 6806 90813 Bibcode2000Natur407908T doi10103835038090 PMID 11057670Chebani Y Marion C Zizzari P Chettab K Pastor M Korostelev M et al April 2016 Enhanced responsiveness of Ghsr Q343X rats to ghrelin results in enhanced adiposity without increased appetite PDF Science Signaling 9 424 ra39 doi101126scisignalaae0374 PMID 27095593 這些對飢餓素反應的神經元對瘦素及胰島素均敏感。Hewson AK Tung LY Connell DW Tookman L Dickson SL December 2002 The rat arcuate nucleus integrates peripheral signals provided by leptin insulin and a ghrelin mimetic Diabetes 51 12 34129 doi102337diabetes51123412 PMID 12453894 而飢餓素會降低胃迷走神經的敏感性，導致較低的胃擴張程度。Page AJ Slattery JA Milte C Laker R ODonnell T Dorian C et al May 2007 Ghrelin selectively reduces mechanosensitivity of upper gastrointestinal vagal afferents American Journal of Physiology Gastrointestinal and Liver Physiology 292 5 G137684 doi101152ajpgi005362006 PMID 17290011 除了其能量平衡的功能，飢餓素還會啟用膽鹼能 多巴胺能犒賞迴路在腹側被蓋區的輸入，以及用來傳達自然犒賞的享樂和增強方面的迴路，Dickson SL Egecioglu E Landgren S Skibicka KP Engel JA Jerlhag E June 2011 The role of the central ghrelin system in reward from food and chemical drugs Molecular and Cellular Endocrinology 340 1 807 doi101016jmce201102017 hdl207726318 PMID 21354264 Whereas ghrelin emerged as a stomachderived hormone involved in energy balance hunger and meal initiation via hypothalamic circuits it now seems clear that it also has a role in motivated rewarddriven behaviours via activation of the socalled cholinergicdopaminergic reward link 如食物和成癮性藥物 Jerlhag E Egecioglu E Dickson SL Andersson M Svensson L Engel JA March 2006 Ghrelin stimulates locomotor activity and accumbal dopamineoverflow via central cholinergic systems in mice implications for its involvement in brain reward Addiction Biology 11 1 4554 doi101111j13691600200600002x PMID 16759336Jerlhag E Egecioglu E Dickson SL Douhan A Svensson L Engel JA March 2007 Ghrelin administration into tegmental areas stimulates locomotor activity and increases extracellular concentration of dopamine in the nucleus accumbens Addiction Biology 12 1 616 doi101111j13691600200600041x PMID 17407492 的中腦邊緣通路。Naleid AM Grace MK Cummings DE Levine AS November 2005 Ghrelin induces feeding in the mesolimbic reward pathway between the ventral tegmental area and the nucleus accumbens Peptides 26 11 22749 doi101016jpeptides200504025 PMID 16137788 該迴路上可以找到飢餓素的受體。而酒精 Jerlhag E Egecioglu E Landgren S Salomé N Heilig M Moechars D et al July 2009 Requirement of central ghrelin signaling for alcohol reward Proceedings of the National Academy of Sciences of the United States of America 106 27 1131823 Bibcode2009PNAS10611318J doi101073pnas0812809106 PMC 2703665 PMID 19564604 及可口 獎勵食物 Egecioglu E Jerlhag E Salomé N Skibicka KP Haage D BohloolyY M et al July 2010 Ghrelin increases intake of rewarding food in rodents Addiction Biology 15 3 30411 doi101111j13691600201000216x PMC 2901520 PMID 20477752Skibicka KP Hansson C Egecioglu E Dickson SL January 2012 Role of ghrelin in food reward impact of ghrelin on sucrose selfadministration and mesolimbic dopamine and acetylcholine receptor gene expression Addiction Biology 17 1 95107 doi101111j13691600201000294x PMC 3298643 PMID 21309956 是需要下視丘飢餓素訊號來提供回饋。 飢餓素與誘導食慾及進食行為有關，而血液迴圈中的飢餓素水平會在飯前最高、飯後最低。Tolle V Bassant MH Zizzari P PoindessousJazat F Tomasetto C Epelbaum J BluetPajot MT April 2002 Ultradian rhythmicity of ghrelin secretion in relation with GH feeding behavior and sleepwake patterns in rats Endocrinology 143 4 135361 doi101210endo14348712 PMID 11897692Cummings DE Frayo RS Marmonier C Aubert R Chapelot D August 2004 Plasma ghrelin levels and hunger scores in humans initiating meals voluntarily without time and foodrelated cues American Journal of Physiology Endocrinology and Metabolism 287 2 E297304 doi101152ajpendo005822003 PMID 15039149 在人和大鼠注射飢餓素已證明會增加食物攝取，即注射量越多，食物攝取越多 Wren AM Small CJ Ward HL Murphy KG Dakin CL Taheri S et al November 2000 The novel hypothalamic peptide ghrelin stimulates food intake and growth hormone secretion Endocrinology 141 11 43258 doi101210endo141117873 PMID 11089570，但飢餓素所增加的不是進食量，而是進食的次數。Faulconbridge LF Cummings DE Kaplan JM Grill HJ September 2003 Hyperphagic effects of brainstem ghrelin administration Diabetes 52 9 22605 doi102337diabetes5292260 PMID 12941764 飢餓素的注射還會增加動物覓食行為，同時增加嗅覺尋找能力，傾向蒐集囤積食物。由於體重是透過能量平衡來調節的，飢餓素濃度與體重會呈負相關，由此飢餓素可以做為肥胖警訊。 血液中濃度 飢餓素在血液中的濃度在 pmol l 的範圍內。具有活性的飢餓素含量和總飢餓素含量都可以被測量出來 Yokota I Kitamura S Hosoda H Kotani Y Kangawa K April 2005 Concentration of the noctanoylated active form of ghrelin in fetal and neonatal circulation Endocrine Journal 52 2 2716 doi101507endocrj52271 PMID 15863960 。迴圈中的飢餓素濃度在進食前升高，並在進食後下降，並且對於蛋白質和碳水化合物的反應比對脂質的反應更強烈。 飢餓素受體 飢餓素受體 GHSR1A（由飢餓素受體剪接而來）可以調節多項飢餓素造成的生物效應，包括：刺激生長激素的釋放、增加飢餓感、調節葡萄糖和脂質的代謝、調節腸胃蠕動和分泌、保護神經和心血管細胞以及調節免疫功能。Yin Y Li Y Zhang W March 2014 The growth hormone secretagogue receptor its intracellular signaling and regulation International Journal of Molecular Sciences 15 3 483755 doi103390ijms15034837 PMC 3975427 PMID 24651458 它們大量存在於下視丘和腦垂體、迷走神經上（在傳入的細胞體和傳出的神經末梢上）以及整個胃腸道中。 作用地點 葡萄糖代謝 整個飢餓素系統（包含 dAG、AG、GHSR、GOAT）皆具有調節葡萄糖的作用。Heppner KM Tong J July 2014 Mechanisms in endocrinology regulation of glucose metabolism by the ghrelin system multiple players and multiple actions European Journal of Endocrinology 171 1 R2132 doi101530EJE140183 PMID 24714083 睡眠 初步研究表明，飢餓素參與了晝夜規律的調節。雖然有文獻表示，沒有發現有力的證據顯示限制睡眠會影響飢餓素以及瘦素的濃度或能量消耗。Zhu B Shi C Park CG Zhao X Reutrakul S June 2019 Effects of sleep restriction on metabolismrelated parameters in healthy adults A comprehensive review and metaanalysis of randomized controlled trials Sleep Medicine Reviews 45 1830 doi101016jsmrv201902002 PMID 30870662 生殖系統 飢餓素對促性腺激素釋放激素（GnRH）的分泌具有抑制作用，可能導致生育力下降。Comninos AN Jayasena CN Dhillo WS 2014 The relationship between gut and adipose hormones and reproduction Human Reproduction Update 20 2 15374 doi101093humupddmt033 PMID 24173881 胎兒和新生兒 飢餓素在胎兒時期的早期肺臟產生，並具有促進肺生長的功能。Santos M Bastos P Gonzaga S Roriz JM Baptista MJ NogueiraSilva C et al April 2006 Ghrelin expression in human and rat fetal lungs and the effect of ghrelin administration in nitrofeninduced congenital diaphragmatic hernia Pediatric Research 59 4 Pt 1 5317 doi10120301pdr000020274866359a9 PMID 16549524 臍帶血中飢餓素的濃度也顯示，飢餓素濃度和新生兒出生體重之間的相關性。 厭食症和肥胖 肥胖個體血漿中的飢餓素濃度低於瘦型個體 Shiiya T Nakazato M Mizuta M Date Y Mondal MS Tanaka M et al January 2002 Plasma ghrelin levels in lean and obese humans and the effect of glucose on ghrelin secretion The Journal of Clinical Endocrinology and Metabolism 87 1 2404 doi101210jcem8718129 PMID 11788653，由此顯示，飢餓素並不直接造成肥胖，只有在小胖威利症候群引起的肥胖中，高濃度的飢餓素能夠造成食物攝入量的增加 Goldstone AP Thomas EL Brynes AE Castroman G Edwards R Ghatei MA et al April 2004 Elevated fasting plasma ghrelin in praderwili syndrome adults is not solely explained by their reduced visceral adiposity and insulin resistance The Journal of Clinical Endocrinology and Metabolism 89 4 171826 doi101210jc2003031118 PMID 15070936。與體重過輕和正常體重的兩個對照組相比，神經性厭食症的血漿飢餓素濃度更高 Misra M Klibanski A July 2014 Endocrine consequences of anorexia nervosa The Lancet Diabetes Endocrinology 2 7 58192 doi101016S22138587 13 701803 PMC 4133106 PMID 24731664Tolle V Kadem M BluetPajot MT Frere D Foulon C Bossu C et al January 2003 Balance in ghrelin and leptin plasma levels in anorexia nervosa patients and constitutionally thin women The Journal of Clinical Endocrinology and Metabolism 88 1 10916 doi101210jc2002020645 PMID 12519838Germain N Galusca B Le Roux CW Bossu C Ghatei MA Lang F et al April 2007 Constitutional thinness and lean anorexia nervosa display opposite concentrations of peptide YY glucagonlike peptide 1 ghrelin and leptin The American Journal of Clinical Nutrition 85 4 96771 doi101093ajcn854967 PMID 17413094。體型較為削瘦的人在一天中，從午夜到黎明的這段時間的飢餓素濃度較高，這顯示肥胖者的晝夜規律系統存在缺陷 Yildiz BO Suchard MA Wong ML McCann SM Licinio J July 2004 Alterations in the dynamics of circulating ghrelin adiponectin and leptin in human obesity Proceedings of the National Academy of Sciences of the United States of America 101 28 104349 Bibcode2004PNAS10110434Y doi101073pnas0403465101 PMC 478601 PMID 15231997。同時，癌症引起的惡體質患者也有高濃度的飢餓素 Garcia JM GarciaTouza M Hijazi RA Taffet G Epner D Mann D et al May 2005 Active ghrelin levels and active to total ghrelin ratio in cancerinduced cachexia The Journal of Clinical Endocrinology and Metabolism 90 5 29206 doi101210jc20041788 PMID 15713718。尚無足夠的證據得出支援或反對使用生長素釋放肽治療癌症相關惡體質的結論 Khatib MN Shankar AH Kirubakaran R Gaidhane A Gaidhane S Simkhada P Quazi Syed Z February 2018 Ghrelin for the management of cachexia associated with cancer The Cochrane Database of Systematic Reviews 2 CD012229 doi10100214651858cd012229pub2 PMC 6491219 PMID 29489032。 相關疾病的控管與治療 胃繞道手術 與體型瘦削和節食減肥的人相比，進行胃繞道手術的人不僅減少了腸道容量，甚至能夠降低體內飢餓素濃度。Cummings DE Weigle DS Frayo RS Breen PA Ma MK Dellinger EP Purnell JQ May 2002 Plasma ghrelin levels after dietinduced weight loss or gastric bypass surgery The New England Journal of Medicine 346 21 162330 doi101056NEJMoa012908 PMID 12023994 在此方面，尚未有研究闡明，接受胃繞道手術的人，體內的飢餓素濃度是否在減輕的體重穩定下來後就恢復正常濃度。Cummings DE Shannon MH July 2003 Ghrelin and gastric bypass is there a hormonal contribution to surgical weight loss The Journal of Clinical Endocrinology and Metabolism 88 7 29993002 doi101210jc2003030705 PMID 12843132 統計也顯示，胃繞道的相關手術中，進行的人從長遠來看，體內飢餓素濃度能夠降低約 60％。Bohdjalian A Langer FB ShakeriLeidenmühler S Gfrerer L Ludvik B Zacherl J Prager G May 2010 Sleeve gastrectomy as sole and definitive bariatric procedure 5year results for weight loss and ghrelin Obesity Surgery 20 5 53540 doi101007s1169500900666 PMID 20094819 參見 參考文獻 Category 位於 3 號人類染色體的基因 Category 生長軸激素 Category 神經肽 Category 肥胖症 Category 肽類激素 Category 世界反興奮劑機構停用物質