Hepatic caecum of amphioxus and origin of vertebrate liver
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1 HEREDITAS (Beijing) , 32(5): ISSN 综述 DOI: /SP.J 文昌鱼肝盲囊与脊椎动物肝脏起源 李红岩, 张士璀, 早期形态和胚胎学研究结果表明, 文昌鱼哈氏窝 内柱和肝盲囊分别是脊椎动物脑垂体 甲状腺和肝脏的同源器官 文章总结了近年来有关文昌鱼肝盲囊与脊椎动物肝脏之间关系的研究成果, 揭示文昌鱼肝盲囊和脊椎动物肝脏具有同源性, 并证明文昌鱼中存在类似脊椎动物的 GH/IGF (Growth hormone/insulin-like growth factor) 和 TH/THR (Thyroid hormone/thyroid hormone receptor) 信号通路, 为脊椎动物肝脏起源于文昌鱼肝盲囊样结构提供了分子水平证据 : 文昌鱼 ; 肝盲囊 ; 肝脏起源 ; GH/IGF; TH/THR Hepatic caecum of amphioxus and origin of vertebrate liver LI Hong-Yan, ZHANG Shi-Cui Department of Marine Biology, Ocean University of China, Qingdao , China Abstract: Previous studies on morphology and embryology revealed that the Hatschek's pit, endostyle and hepatic caecum in amphioxus were homologous to the precursors of pituitary, thyroid and liver in vertebrates, respectively. Here, we summarize the recent advances on the relationship between amphioxus hepatic caecum and the vertebrate liver as well as the origin of growth hormone/insulin-like growth factor (GH/IGF) and thyroid hormone/thyroid hormone receptor (TH/THR) signal pathways, which will provide molecular support to the homology of amphioxus hepatic caecum to vertebrate liver. Keywords: amphioxus; hepatic caecum; origin of liver; GH/IGF; TH/THR, 1886, Kowalevsky ( ), ( ),, (missing link),, Kowalevsky, ( ) ( ), 5.5, [1],, ( ) [2, 3], ( 1), 收稿日期 : ; 修回日期 : 基金项目 : ( , , ) 作者简介 : (1975 ),,, hongyanli@ouc.edu.cn 通讯作者 : (1957 ),,,, sczhang@ouc.edu.cn
2 438 HEREDITAS (Beijing) 图 1 脊索动物演化关系 ( ) ( ) ( ) a b 2, 35,,,, 1, ( ) 2,,,,,, Hammar [4] 1898,,, :,,,, Hammar, Barrington [5], 1975, Welsch [6],,,,, (Vg) (AAT) (AT) (Pg) Han [7] Vg Liang [8, 9] Lun [10] AAT AT Pg Pg, AAT AT Pg,, S- (GST) B [11, 12], GST B, GST B
3 5 : 439 2,, 2 GH/IGF (Growth hormone, GH),,, GH GH, (Insulin-like growth factor, IGF) IGF (Prepro-insulin), IGF -, GH/IGF GH IGF, IGF IGF (IGF binding proteins, IGFBPs) IGFBPs,,,, (Hatschek s pit) [13], 80,, (Luteinizing hormone, LH) (Follicle-stimulating hormone, FSH) [14] (Gonadotrophin-releasing hormone A, GnRH-A) [15] [16] - β- (Human luteinizing hormone β, hlhβ) (Human chorionic gonadotrophin, hcg) [17] - (Gonadotropic hormone, GTH) [18],,, - GH/IGF, Guo [19] IGF, IGF, IGF, IGF, GHR GH/IGF ( 2),, GH, GH 3 TH/THR (Thyroid hormones; THs) T 4 T 3, THs, THs,, [20] THs [21~23], THs, THs (THRs) THRs, THRα THRβ THs THRs TH-THR,, T 3 C/EBP(CCAAT/enhancerbinding proteins) C/EBPα C/EBPβ [24~27] [28, 29] THs TH/THR, 1873, Müller , [30, :, T 3 T 31] 4 [32, 33] [5, 34]
4 440 HEREDITAS (Beijing) 图 2 GH/IGF 和 TH/THR 信号通路, GH, GHR, IGF, GH/IGF THS, TH, THR, C/EBP, TH/THR, GH IGF, GH/IGF TH, C/EBP, TH/THR Pax-8 Nkx2-1,,, Nkx2-1 AmphiNk2-1 Pax-2 Pax-5 Pax-8 AmphiPax-2/5/8 AmphiNk2-1,, 7 d, AmphiNk2-1 [35] AmphiPax-2/5/8,, AmphiPax-2/5/8, 7 d,, AmphiNk2-1 AmphiPax-2/5/8,, TH/THR, Wang [36] TH THR, T 4 T 3 TRIAC C/EBPα/β THR T 3 TRIAC TH/THR ( 2), THs THs (Thyroidstimulating hormone, TSH), THs,
5 5 : :,,, GH/IGF TH/THR, 2,,, 2, 参考文献 (References): [1] Stach T. Chordate phylogeny and evolution: a not so simple three-taxon problem. J Zool, 2008, 276(2): [2] Delsuc F, Brinkmann H, Chourrout D, Philippe H. Tunicates and not cephalochordates are the closest living relatives of vertebrates. Nature, 2006, 439(7079): [3] Putnam NH, Butts T, Ferrier DE, Furlong RF, Hellsten U, Kawashima T, Robinson-Rechavi M, Shoguchi E, Terry A, Yu JK, Benito-Gutiérrez EL, Dubchak I, Garcia-Fernàndez J, Gibson-Brown JJ, Grigoriev IV, Horton AC, de Jong PJ, Jurka J, Kapitonov VV, Kohara Y, Kuroki Y, Lindquist E, Lucas S, Osoegawa K, Pennacchio LA, Salamov AA, Satou Y, Sauka-Spengler T, Schmutz J, Shin-I T, Toyoda A, Bronner-Fraser M, Fujiyama A, Holland LZ, Holland PW, Satoh N, Rokhsar DS. The amphioxus genome and the evolution of the chordate karyotype. Nature, 2008, 453(7198): [4] Hammar JA. Zur Kenntnis der Leberentwichlung bei Amphioxus. Anat Anz, 1898, 14, [5] Barrington EJW. The digestive system of Amphioxus (Branchiostoma lancelatus). Phil Trans Roy Soc Lond series B, 1937, 228: [6] Welsch U. The fine structure of the pharynx, crytopodocytes and digestive caecum of amphioxus (Branchiostoma lanceolatum). Symp Zool Soc Lond, 1975, 36: [7] Han L, Zhang SC, Wang YJ, Sun XT. Immunohistochemical localization of vitellogenin in the hepatic diverticulum of the amphioxus Branchiostoma belcheri tsingtauense, with implications of the origin of the liver. Invert Biol, 2006, 125(2): [8] Liang YJ, Zhang SC, Lun LM, Han L. Presence and localization of antithrombin and its regulation after acute lipopolyssacharide exposure in amphioxus, with implications for the origin of vertebrate liver. Cell Tissue Res, 2006, 323(3): [9] Liang YJ, Zhang SC. Demonstration of plasminogen-like protein in amphioxus with implications of the origin of vertebrate liver. Acta Zool (Stockholm), 2006, 87(2): [10] Lun LM, Zhang SC, Liang YJ. Alanine aminotransferase in amphioxus: presence, localization and up-regulation after acute lipopolysaccharide exposure. J Biochem Mol Biol, 2006, 39(5): [11] Fan CX, Zhang SC, Liu ZH, Li L, Luan J, Saren G. Identification and expression of a novel class of glutathione-s-transferase from amphioxus Branchiostoma belcheri with implications to the origin of vertebrate liver. Int J Biochem Cell Biol, 2007, 39(2): [12] He YN, Tang B, Zhang SC, Liu ZH, Zhao B, Chen LL. Molecular and immunochemical demonstration of a novel member of Bf/C2 homolog in amphioxus Branchiostoma belcheri: implications for involvement of hepatic cecum in acute phase response. Fish Shellfish Immunol, 2008, 24(6): [13] Tjoy LT, Welsch U. Electron microscopical observations on Kolliker s and Hatschek s pit on the wheel organ in the head region of amphioxus (Branchiostoma lanceolatum). Cell Tissue Res, 1974, 153(2): [14] Chang CY, Liu YH, Zhu HH. The sex steroid hormones and their functional regulation in amphioxus (Branchiostoma belcheri Gray). Proc 9 th Internat Symp Comp Endocrinol, 1981, 9. [15].., 1998, 43(3): [16],.., 1984, 17(1): [17],,. (LH)., 1982, 27(15): [18].
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