中国乳业 ›› 2021, Vol. 0 ›› Issue (9): 21-31.doi: 10.12377/1671-4393.21.09.04

• 尿素氮营养专题 • 上一篇    下一篇

反刍动物体内尿素循环及其转运蛋白的分子调控机制研究进展

甘水燕1, 刘虎1, 周建伟1,*   

  1. 草地农业生态系统国家重点实验室; 兰州大学草地农业科技学院,甘肃兰州 730020
  • 出版日期:2021-09-25 发布日期:2021-09-29
  • 通讯作者: *周建伟(1985-),男,浙江兰溪人,研究员,硕士生导师,研究方向为蔵系反刍家畜营养代谢与调控。
  • 作者简介:甘水燕(1998-),女,广西贵港人,硕士,研究方向为反刍动物营养代谢与调控;刘 虎(1992-),男,湖南益阳人,博士研究生,研究方向为藏系反刍动物营养代谢与生态。
  • 基金资助:
    甘肃省重点研发计划项目(21YF5WA117)

Progresses of Research on Urea Recycling and Molecular Regulation Mechanism of Urea Transportersin Ruminants

GAN Shuiyan1, LIU Hu1, ZHOU Jianwei1,*   

  1. State Key Laboratory of Grassland Agro-Ecosystems,College of Pastoral Agriculture Science and Technology,Lanzhou University,Lanzhou Gansu 730020
  • Online:2021-09-25 Published:2021-09-29

摘要: 虽然在哺乳动物体内都存在尿素循环利用机制,但是反刍动物由于瘤胃的存在而使得尿素循环在维系机体氮平衡和提高氮素利用效率等方面发挥着更加重要的生物学意义。通过瘤胃壁扩散或转运,血液中的尿素可进入胃肠道,在脲酶的作用下转化为氨态氮,从而为瘤胃微生物蛋白合成提供氮源。研究表明,尿素在瘤胃上皮的自由扩散速度较慢,而尿素转运蛋白可以介导尿素分子高效地进行跨膜转运,其也被认为是反刍动物尿素循环最重要的调控因子。然而,相关报道已经证实,尿素转运蛋白的表达和功能发挥受到日粮营养水平与结构组成、瘤胃内环境、动物年龄等因素的影响。本文以尿素循环为出发点,重点阐述了反刍动物体内尿素循环的特点、影响因素以及尿素转运蛋白的表达特征及其分子调控过程,以期从分子生物学角度解析反刍动物尿素循环的作用机制,从而为生产实践中动物氮素的精准营养提供理论依据和技术支撑。

关键词: 尿素循环, 尿素转运蛋白, 水通道蛋白, 反刍动物

Abstract: The mechanism of urea recycling is existing in all mammals. However,it plays a crucially biological role in ruminants for maintaining nitrogen balance and improving the efficiency of nitrogen utilization because of the urea transportation in rumen. This specific mechanism in ruminants is able to make blood urea from urinary excretionrepartitioned into the gastrointestinal tract by the free diffusion or transfer throughout the rumen epithelium. Subsequently, the urea would be converted to ammonia nitrogen upon the reaction of urease, which then provides nitrogen source for rumen microbial protein synthesis. Many previous studies have shown that it was rather slow of the speed of urea transferred across the rumen epithelium by free diffusion. However, urea transporters are known to be involved in urea transferring across rumen epithelium and thus facilitate efficiently the urea nitrogen recycling process. Consequently, urea transporters are considered as the most important regulators of the urea recycling mechanism in ruminants. Nevertheless,relevant publications have demonstrated that the expression and function characteristics of urea transporters were regulated by numerous factors,such as dietary nutrients content and structural constitutes,intraluminal environment,and the age of animals. Based on the above backgrounds,this paper was conducted to overview the characteristics of urea recycling in ruminants,and also for the introduction of the effect factors,expression characteristics and the molecular regulation of urea transporters. It is therefore,the present study was aimed to interpret the mechanism of urea recycling in ruminants from the perspective of molecular biology,and then to provide theoretical basis and technical support for the precise nitrogen nutrition of ruminant production practice.

Key words: urea recycling, urea transporters, aquaglyceroporins, ruminant

[1] Marini J C,Klein J D,Sands J M,et al.Effect of nitrogen intake on nitrogen recycling and urea transporter abundance in lambs[J]. Journal of Animal Science,2004,82(4):1157-1164.
[2] Leip A,Billen G,Garnier J,et al.Impacts of European livestock production:nitrogen,sulphur,phosphorus and greenhouse gas emissions,land-use,water eutrophication and biodiversity[J]. Environment Research Letters,2015,10(11):1-13.
[3] 高民,温雅俐,杜瑞平. 反刍动物生产中碳、氮、磷、钾的减排问题[J].畜牧与饲料科学,2011,32(Z1):94-95,99.
[4] 徐红蕊,时建青,赵国琦. 反刍动物体内尿素氮代谢研究进展[J]. 畜牧兽医杂志,2006,25(1):23-27.
[5] Stewart G S,Graham C,Cattell S, et al.UT-B is expressed in bovine rumen: potential role in ruminal urea transport[J]. American Journal of Physiology-Regulatory, Integrative and Comparative Physiology,2005,289(2):605-612.
[6] Walpole M E,Schurmann B L,Górka P, et al.Serosal-to-mucosal urea flux across the isolated ruminal epithelium is mediated via urea transporter-B and aquaporins when Holstein calves are abruptly changed to a moderately fermentable diet[J]. Journal of Dairy Science. 2015,98(2):1204-1213.
[7] Rojen B A,Poulsen S B,TheilP K,et al.Effects of dietary nitrogen concentration onmessenger RNA expression and protein abundance of urea transporter B and aquaporins in ruminalpapillae from lactating Holstein cows[J]. Journal of Dairy Science,2011, 94(5):2587-2591.
[8] Stewart G S,Smith C P.Urea nitrogen salvage mechanisms and their relevance to ruminants,non-ruminants and man[J]. Nutrition Research Reviews,2005,18(1):49-62.
[9] Obara Y,Dellow D W,Nolan J V.The influence of energy-rich supplements on nitrogen kinetics in ruminants[M]. Physiological Aspects of Digestion & Metabolism in Ruminants,Academic Press,1991:515-539.
[10] Harmeyer J, Martens H.Aspects of urea metabolism in ruminants with reference to the goat[J]. Journal of Dairy Science,1980, 63(10):1707-1728.
[11] 周建伟. 藏羊对青藏高原氮素营养胁迫的适应性研究:[博士论文][D]. 兰州大学,2015.
[12] Doranalli K,Penner G B,Mutsvangwa T.Feeding oscillating dietary crude protein concentrations increases nitrogen utilization in growing lambs and this response is partly attributable to increased urea transfer to the rumen[J]. The Journal of Nutrition,2011, 141(4):560-567.
[13] Müller C B M,Görs S,Derno M,et al. Differences between Holstein dairy cows in renal clearance rate of urea affect milk urea concentration and the relationship between milk urea and urinary nitrogen excretion[J]. Science of The Total Environment,2020, 755(Pt 2).
[14] Cécile C,Pierre P,Germain.R.Cloning and functional characterization of a rat urea transporter: expression in the brain[J]. Biochimica et Biophysica Acta (BBA)-Gene Structure and Expression,1996,1309(3):197-199.
[15] Fenton R A,Shodeinde A,Knepper M A.UT-A urea transporter promoter,UT-Aα,targets principal cells of the renal inner medullary collecting duct[J]. American Journal of Physiology Renal Physiology, 2006,290(1):188-195.
[16] Levin E J,Cao Y,Enkavi G,et al.Mechanism and regulation of urea permeation in a mammalian urea channel[J]. Biophysical Journal,2012,102(3):215.
[17] Levin E J,Ming Z.Structure of urea transporters[J]. Sub-Cellular Biochemistry,2014,73:65-78.
[18] Lucien N,Bruneval P,Lasbennes F, et al.UT-B1 urea transporter is expressed along the urinary and gastrointestinal tracts of the mouse[J]. American Journal of Physiology Regulatory Integrative & Comparative Physiology,2005,288(4):1046-1056.
[19] Collins D,Winter D C,Hogan A M,et al.Differential protein abundance and function of UT-B urea transporters in human colon[J]. American Journal of Physiology,2010,298(1):345-351.
[20] Marie M T T T,Jean-P C,Lise B. Molecular basis for the dialysis disequilibrium syndrome:altered aquaporin and urea transporter expression in the brain[J]. Nephrology,Dialysis,Transplantation:Official Publication of the European Dialysis and Transplant Association-European Renal Association,2005,20(9):1984-1988.
[21] Tickle P,Thistlethwaite A,Smith C P,et al.Novel bUT-B2 urea transporter isoform is constitutively activated[J]. American Journal of Physiology Regulatory Integrative & Comparative Physiology,2009,297(2):323-329.
[22] Simmons N L,Chaudhry A S,Graham C,et al.Dietary regulation of ruminal bovine UT-B urea transporter expression and localization[J]. Journal of Animal Science, 2009, 87(10):3288-3299.
[23] 仲崇亮. 尿素转运蛋白UT-B和AQP3在瘤胃的表达、分布和调节机制:[博士论文][D].兰州大学,2021.
[24] Olives B,Merriman M,Bailly P,et al.The molecular basis of the kidd blood group polymorphism and its lack of association with type 1 diabetes susceptibility[J]. Human Molecular Geneticst,1997,6(7):1017-1020.
[25] Promeneur D,Rousselet G,Bankir L,et al.Evidence for distinct vascular and tubular urea transporters in the rat kidney[J]. Journal of the American Society of Nephrology,1996,7(6):852-860.
[26] Bradford A D,Terris J,Ecelbarger CA,et al.97 and 117 kDa forms of the collecting duct urea transporter UT-A1 are due to different states of glycosylation[J]. American Journal of Physiology-Renal Physiology,2001,281(1):133-143.
[27] Nielsen S,Terris J,Smith CP,et al.Cellular and subcellular localization of thevasopressin-regulated urea transporter in rat kidney[J]. Proceedings of the National Academy of Sciences of the United States of America,1996,93(11):5495-5500.
[28] Couriaud C, Ripoche P, Rousselet G.Cloning and functional characterization of a rat urea transporter: expression in the brain[J]. Biochimicaet Biophysica Acta(BBA)-Gene Structure Expression,1996, 1309(3):197-199.
[29] Timmer R T,Klein J D,Bagnasco SM,et al.Localization of the urea transporter UT-B protein in human and rat erythrocytes and tissues[J]. American Journal of Physiology Cell Physiology,2001,281(4):1318-1325.
[30] Bagnasco SM,Peng T,Janech M G,et al.Cloning and characterization of the human urea transporter UT-A1 and mapping of the human Slc14a2 gene[J]. American Journal of Physiology-Renal Physiology,2001,281(3):400-406.
[31] Smith C P,Lee W S,Martial S,et al.Cloning and regulation of expression of the rat kidney urea transporter (rUT2)[J]. Journal of Clinical Investigation,1995,96(3):1556-1563.
[32] Doran J J,Timmer R T,Sands J M.Accurate mRNA size determination in northern analysis using individual lane size markers[J]. Biotechniques,1999,27(2):280-282.
[33] Potter E A,Stewart G,Smith C P.Urea flux across MDCK-mUT-A2 monolayers is acutely sensitive to AVP,cAMP,and [Ca2+]i[J]. American Journal of Physiology-Renal Physiology,2006,291(1):122-128.
[34] Terris J M,Knepper M A,Wade J B.UT-A3:localization and characterization of an additional urea transporter isoform in the IMCD[J]. American Journal of Physiology-Renal Physiology, 2001,280(2):325-332.
[35] Karakashian A,Timmer R T,Klein JD,et al.Cloning and characterization of two new mRNA isoforms of the rat renal urea transporter:UT-A3 and UT-A4[J]. Journal of the American Society of Nephrology,1999,10(2):230-237.
[36] Fenton R A,Howorth A,Cooper G J,et al.Molecular characterization of a novel UT-A urea transporter isoform (UT-A5) in testis[J]. American Journal of Physiology-Cell Physiology,2000,279(5):1425-1431.
[37] Echevarria M,Windhager E E,Frindt T G.Cloning and expression of AQP3,a water channel from the medullary collecting duct of rat kidney[J]. Proceedings of the National Academy of Sciences of the United States of America, 1994,91(23):10997-11001.
[38] Ishibashi K,Kuwahara M, Yong G,et al.Cloning and functional expression of a new aquaporin (AQP9) abundantly expressed in the peripheral leukocytes permeable to water and urea, but not to glycerol[J]. Biochemical & Biophysical Research Communications,1998,244(1):268-274.
[39] Ishibashi K,Morinaga T,Kuwahara M,et al.Cloning and identification of a new member of water channel (AQP10)as an aquaglyceroporin[J]. Biochimicaet Biophysica Acta(BBA)-Gene Structure Expression,2002,1576(3):335-340.
[40] Li C,Wang W.Urea transport mediated by aquaporin water channel proteins[J]. Sub-Cellular Biochemistry,2014,73:227-265.
[41] And J,Agre P.Hourglass pore-forming domains restrict aquaporin-1 tetramer assembly-biochemistry (ACS publications)[J]. Biochemistry,1999,38(3):923-9288.
[42] Scott K A,Penner G B, Timothy M.Influence of forage level and corn grain processing on whole-body urea kinetics, and serosal-to-mucosal urea flux and expression of urea transporters and aquaporins in the ovine ruminal, duodenal, and cecal epithelia[J]. Journal of Animal Science, 2020,98(4):1-16.
[43] Zhong C,Farrell A,Stewart G S.Localization of aquaporin-3 proteins in the bovine rumen[J]. Journal of Dairy Science,2020, 103( 3):2814-2820.
[44] Lei T,Zhou L,Layton A T,et al.Role of thin descending limb urea transport in renal urea handling and the urine concentrating mechanism[J]. American Journal of Physiology Renal Physiology, 2011,301(6):1251-1259.
[45] Fenton R A,Yang B.Urea transporter knockout mice and their renal phenotypes[J]. Sub-Cellular Biochemistry,2014,73:137-152.
[46] Starke S,Muscher A S,Hirschhausen N,et al.Expression of urea transporters is affected by dietary nitrogen restriction in goat kidney[J]. Journal of Animal Science,2012,90(11):3889-3897.
[47] 康婧鹏. 饲粮能量水平对藏羊尿素循环及其组织中UT-B表达量的影响:[硕士论文][D].兰州大学,2018.
[48] 郭亚敏. 饲粮能量水平对藏羊瘤胃氮素转运及利用效率的影响:[硕士论文][D].兰州大学,2019.
[49] 张莹,郭旭生,丁路明,等.反刍动物尿素氮再循环利用的研究进展[J].动物营养学报,2009,21(5):609-616.
[50] 樊艳华,孙海洲,桑丹,等.提高反刍动物氮利用率的研究进展[J].家畜生态学报,2014,35(10):1-9.
[51] Van Amburgh M E,Overton T R,Chase L E,et al. The cornell net carbohydrate and protein system:current and future approaches for balancing of amino acids[C]//Proceedings of Cornell Nutrition Conference for Feed Manufacturers, Syracuse, NY. New York State College of Agriculture & Life Sciences, Cornell University,2009.
[52] Marini J C, Van Amburgh M E. Nitrogen metabolism and recycling in Holstein heifers[J]. Journal of Animal Science, 2003, 81(2):545-552.
[53] Abadi E,Tahmasebi A M, MD Mesgaran, et al.Effect of dietary crude protein level on UT-B expression and nitrogen efficiency in growing baluchi male lambs fed low or high concentrate diets[J]. Iranian Journal of Applied Animal Science,2015,5(2):323-332.
[54] 艳城. 日粮对细毛羊瘤胃上皮的SCFA吸收相关基因及氮素转运的调控:[硕士论文][D].内蒙古农业大学,2015.
[55] Elena S, Mirco C, Francesco G, et al.Effect of dietary nitrogen level and source on mRNA expression of urea transporters in the rumen epithelium of fattening bulls[J]. Archives of Animal Nutrition,2018,72(5):341-350.
[56] Minuti A, Palladino A, Khan M J,et al.Abundance of ruminal bacteria, epithelial gene expression, and systemic biomarkers of metabolism and inflammation are altered during the peripartal period in dairy cows[J]. Journal of Dairy Science, 2015, 98(12):8940-8951.
[57] Berends H,van den Borne J J G C,Rojen B A,et al. Urea recycling contributes to nitrogen retention in calves fed milk replacer and low-protein solid feed[J]. Journal of Nutrition,2014,144(7):1043.
[58] 王笑笑. 碳水化合物来源对泌乳奶牛氮素利用、尿液代谢组与瘤胃上皮细胞尿素转运蛋白和水通道蛋白表达的影响:[博士论文][D]. 河南农业大学, 2016.
[59] Lu Z, Gui H, Lei Y, et al.Short-chain fatty acids and acidic pH upregulate UT-B, GPR41, and GPR4 in rumen epithelial cells of goats[J]. American Journal of Physiology Regulatory Integrative & Comparative Physiology,2015,308(4):283-293.
[60] De Oliveira C V R, Silva T E, Batista E D, et al. Urea supplementation in rumen and post-rumen for cattle fed a low-quality tropical forage[J]. British Journal of Nutrition,2020,124(11):1166-1178.
[61] Lu Z,Stumpff,Deiner C,et al.Modulation of sheep ruminal urea transport by ammonia and pH[J]. American Journal of Physiology&Regulatory,2014,307(5):558-570.
[1] 陆钟岩, 张雯萱, 阿合拉·托留拜. 日粮添加尿素对瘤胃上皮细胞增殖、凋亡以及吸收转运能力的影响[J]. 中国乳业, 2021, 0(9): 8-20.
[2] 王梦芝, 杨斯涵, 刘福元, 张振斌, 赵建. 缓释尿素制备工艺及其在反刍动物生产中应用的研究进展[J]. 中国乳业, 2021, 0(9): 32-39.
[3] 熊展博, 赵圣国, 王加启. 细菌脲酶分解尿素机制及其调控[J]. 中国乳业, 2021, 0(9): 3-7.
[4] 张书阅, 熊本海, 刘明, 蒋林树. 酿酒酵母培养物对瘤胃内环境和免疫功能的影响及其在反刍动物上的应用[J]. 中国乳业, 2021, 0(7): 18-24.
[5] 付瑶, 王俊, 齐志国, 郭江鹏. 高锌日粮对反刍动物的影响及在生产中的应用[J]. 中国乳业, 2019, 0(6): 38-40.
[6] 王国艮, 葛旭升. 过瘤胃蛋白质保护技术在反刍动物饲料中的应用[J]. 中国乳业, 2019, 0(6): 41-46.
[7] 卢昌文, 彭华, 陈雅坤, 梁春宇, 赵连生. 影响青贮品质的流程分析[J]. 中国乳业, 2019, 0(4): 48-52.
[8] 李志春, 闫益波. 奶牛瘤胃酸中毒及其营养调控[J]. 中国乳业, 2019, 0(2): 48-54.
Viewed
Full text


Abstract

Cited

  Shared   
  Discussed   
No Suggested Reading articles found!