水稻抗性淀粉的分子基础和育种研究进展
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摘要
淀粉是人类能量的主要来源,淀粉在消化道中被水解成葡萄糖,进而用于细胞的代谢。当细胞的能量需求减少时,多余的能量作为糖原或脂肪存储在体内。因此,长期的过度饮食和缺乏运动都可能会导致一些健康问题,如肥胖和2型糖尿病。而抗性淀粉(resistant starch, RS)在体内消化缓慢,具有降低血糖和血脂、控制体重、促进矿物质的消化和吸收等多种生理功能,是近年来功能食品领域的研究热点。水稻是人类的重要粮食,但大多数水稻品种的RS含量往往低于3%。因此,开展水稻RS的理论研究和高RS水稻品种的选育非常重要。本综述总结了RS的功能和类型,着重对水稻RS相关基因和分子机理的研究进行了总结,并归纳了中国高RS水稻育种的进展,以期为水稻RS研究和分子育种提供参考。
Starch is the main source of human energy. Starch is normally hydrolyzed into glucose in the digestive tract and further used for metabolism by cells. When the required energy of cells decreases, the excess calories are stored in the body as glycogen or fats. Therefore, long-term overeating and lack of exercises would cause health problems, such as obesity and type 2 diabetes. However, resistant starch(RS) is digested slowly in body, which is helpful to several physiological functions, such as lowering blood glucose and lipid levels, controlling weight,promoting mineral digestion and absorption in human body. Therefore, RS has become a research focus in the field of functional food. Rice is an important food for mankind, but the content of RS in most rice varieties is usually less than 3%. So, it is important to carry out the theoretical research of rice RS and breeding of high RS in rice. The review summarized the functions and types of RS, especially the study of RS-related genes and molecular mechanism of rice. In addition, we introduce the progress in high RS rice breeding in China, so as to provide valuable information and reference for RS study and molecular breeding in rice.
Starch is the main source of human energy. Starch is normally hydrolyzed into glucose in the digestive tract and further used for metabolism by cells. When the required energy of cells decreases, the excess calories are stored in the body as glycogen or fats. Therefore, long-term overeating and lack of exercises would cause health problems, such as obesity and type 2 diabetes. However, resistant starch(RS) is digested slowly in body, which is helpful to several physiological functions, such as lowering blood glucose and lipid levels, controlling weight,promoting mineral digestion and absorption in human body. Therefore, RS has become a research focus in the field of functional food. Rice is an important food for mankind, but the content of RS in most rice varieties is usually less than 3%. So, it is important to carry out the theoretical research of rice RS and breeding of high RS in rice. The review summarized the functions and types of RS, especially the study of RS-related genes and molecular mechanism of rice. In addition, we introduce the progress in high RS rice breeding in China, so as to provide valuable information and reference for RS study and molecular breeding in rice.
引文
A huja G.,Jaiswal S.,and Chibba r R.N.,2013,Resistant starch sources,applications and health benefits,Sources,Applications and Health Benefits,28(46):131-144
Ai Y.,Hasjim J.,and Jane J.L.,2013,Effects of lipids on enzymatic hydrolysis and physical properties of starch,Carbohydrate Polymers,92(1):120-127
Baghurst P.A.,Baghurst K.I.,and Record S.J.,1996,Dietary fibre,non-starch polysaccharides and resistant starch:a review,Food Australia,48(3):S3-S35
Ball S.,Guan H.P.,James M.,Myers A.,Keeling P.,Mouille G.Buléon A.,Colonna P.,and Preiss J.,1996,From glycogen to amylopectin:a model for the biogenesis of the plant starch granule,Cell,86(3):349-352
Bao J.S.,Zhou X.,Xu F.F.,He Q.,and Park Y.J.,2017,Genomewide association study of the resistant starch content in rice grains,Starch/St覿rke,69(7-8):1600343
Bird A.R.,Brown I.L.,and Topping D.L.,2000,Starches,resistant starches,the gut microflora and human health,Curr.Issues in Intest.Microbiol.,1(1):25-37
Brouns F.,Kettlitz B.,and Arrigoni E.,2002,Resistant starch and"the butyrate revolution",Trends in Food Science&Technology,13(8):251-261
Chen L.,Magliano D.J.,and Zimmet P.Z.,2011,The worldwide epidemiology of type 2 diabetes mellitus-present and future perspectives,Nat.Rev.Endocrinol.,8(4):228-236
Dupuis J.H.,Liu Q.,and Yada R.Y.,2015,Methodologies for increasing the resistant starch content of food starches:a review,Comprehensive Reviews in Food Science&Food Safety,13(6):1219-1234
Englyst H.,Wiggins H.S.,and Cummings J.H.,1982,Determination of the non-starch polysaccharides in plant foods by gas-liquid chromatography of constituent sugars as alditol acetates,Analyst,107(1272):307-318
Fitzgerald M.A.,Rahman S.,Resurreccion A.P.,Concepcion J.,Daygon V.D.,Dipti S.S.,Kabir K.A.,Klingner B.,Morell M.K.,and Bird A.R.,2011,Identification of a major genetic determinant of glycaemic index in rice,Rice,4(2):66-74
Guo C.F.,Li J.Y.,and Zhang S.W.,2006,The research development on the physiological function of resistant starch,Shipin Keji(Food Science and Technology),31(2):1-3(郭春锋,李婧妍,张守文,2006,抗性淀粉生理功能研究进展,食品科技,31(2):1-3)
Han J.A.,and Bemiller J.N.,2007,Preparation and physical characteristics of slowly digesting modified food starches,Carbohydrate Polymers,67(3):366-374
Hannah L.C.,and James M.,2008,The complexities of starch biosynthesis in cereal endosperms,Curr.Opin.Biotechnol.,19(2):160-165
Hennen-Bierwagen T.A.,Lin Q.,Grimaud F.,Planchot V.,Keeling P.L.,James M.G.,and Myers A.M.,2009,Proteins from multiple metabolic pathways associate with starch biosynthetic enzymes in high molecular weight complexes:a model for regulation of carbon allocation in maize amyloplasts,Plant Physiol.,149(3):1541-1559
Hu P.,Zhao H.J.,Duan Z.Y.,Zhang L.L.,and Wu D.X.,2004,Starch digestibility and the estimated glycemic score of different types of rice differing in amylose contents,Journal of Cereal Science,40(3):231-237
James M.G.,Denyer K.,and Myers A.M.,2003,Starch synthesis in the cereal endosperm,Curr.Opin.Plant Biol.,6(3):215-222
Kang H.G.,Park S.H.,Matsuoka M.,and An G.H.,2005White-core endosperm floury endosperm-4 in rice is generated by knockout mutations in the C4-type pyruvate orthophosphate dikinase gene(OsPPDKB),Plant J.,42(6):901-911
Luo X.,Huang J.F.,Zhu Y.S.,Xie H.G.,Wu F.X.,Zhang M.Q.Zhang J.F.,and Xie H.A.,2014,Genetic analysis of high resistant starch characteristics for rice variety gongmi 3(Oryza sativa ssp.indica),Nongye Shengwu Jishu Xuebao(Journa of Agricultural Biotechnology),22(1):10-16(罗曦,黄锦峰朱永生,谢鸿光,吴方喜,张木清,张建福,谢华安,2014水稻功米3号高抗性淀粉性状的遗传分析,农业生物技术学报,22(1):10-16)
Myers A.M.,Morell M.K.,James M.G.,and Ball S.G.,2000,Recent progress toward understanding biosynthesis of the amylopectin crystal,Plant Physiol.,122(4):989-997
Nakamura Y.,2002,Towards a better understanding of the metabolic system for amylopectin biosynthesis in plants:rice endosperm as a model tissue,Plant&Cell Physiology,43(7):718-725
Ordonio R.L.,and Matsuoka M.,2016,Increasing resistant starch content in rice for better consumer health,Proc.Natl.Acad Sci.USA,113(45):12616-12618
Raben A.,Tagliabue A.,Christensen N.J.,Madsen J.,Holst J.J.and Astrup A.,1994,Resistant starch:the effect on postprandial glycemia,hormonal response,and satiety,American Journal of Clinical Nutrition,60(4):544-551
Raigond P.,Ezekiel R.,and Raigond B.,2015,Resistant starch in food:a review,J.Sci.Food Agric.,95(10):1968-1978
Sun Z.M.,Bai J.J.,Zhu H.M.,Sun C.L.,Pu Z.Z.,and Hao Z.S.2012,Effects of mixing ratio and processing methods on resistant starch content of products of rice'Jiangtangdao 1'Henongxue Bao(Journal of Nuclear Agricultural Sciences)26(2):318-323(孙志敏,白建江,朱辉明,孙春龙,朴钟泽郝再彬,2012,混配比例及加工方式对“降糖稻1号”稻米产品抗性淀粉含量的影响,核农学报,26(2):318-323)
Tian Z.X.,Qian Q.,Liu Q.Q.,Yan M.X.,Liu X.F.,Yan C.J.,Liu G.F.,Gao Z.Y.,Tang S.Z.,and Zeng D.L.,2009,Allelic diversities in rice starch biosynthesis lead to a diverse array of rice eating and cooking qualities,Proc.Natl.Acad.Sci.US-A,106(51):21760-21765
Tran N.A.,Daygon V.D.,Resurreccion A.P.,Cuevas R.P.,Corpuz H.M.,and Fitzgerald M.A.,2011,A single nucleotide polymorphism in the Waxy gene explains a significant component of gel consistency,Theor.Appl.Genet.,123(4)519-525
Van de Wal M.,D'Hulst C.,Vincken J.P.,Buléon A.,Visser R.,and Ball S.,1998,Amylose is synthesized in vitro by extension of and cleavage from amylopectin,J.Biol.Chem.,273(35):22232-22240
Witt T.,Gidley M.J.,and Gilbert R.G.,2010,Starch digestion mechanistic information from the time evolution of molecular size distributions,J.Agric.Food Chem.,58(14):8444-8452
Wu W.,Liu X.,Yang C.Z.,Shu X.L.,and Wu D.X.,2006,Progresses in the study of resistant starch and functional rice for prevention of diabetes and hyperlipidemia,Henongxue Bao(Journal of Nuclear Agricultural Sciences),20(1):60-63(吴伟,刘鑫,杨朝柱,舒小丽,吴殿星,2006,抗性淀粉及预防糖尿病和肥胖症功能稻米研究进展,核农学报,20(1):60-63)
Yamada Y.,Hosoya S.,Nishimura S.,Tanaka T.,Kajimoto Y.,Nishimura A.,and Kajimoto O.,2005,Effect of bread containing resistant starch on postprandial blood glucose levels in humans,Biosci.Biotechnol.Biochem.,69(3):559-566
Yang C.Z.,Li C.S.,Shu X.L.,Zhang Z.Z.,Zhang L.,Zhao H.J.,Ma C.X.,and Wu D.X.,2005,Starch properties of rice mutant enriched with resistant starch,Zhongguo Shuidao Kexue(Chinese Journal of Rice Science),19(6):516-520(杨朝柱,李春寿,舒小丽,张志转,张磊,赵海军,马传喜,吴殿星,2005,富含抗性淀粉水稻突变体的淀粉特性,中国水稻科学,19(6):516-520)
Yang C.Z.,Shu X.L.,Zhang L.L.,Wang X.Y.,Zhao H.J.,Ma C.X.,and Wu D.X.,2006,Starch properties of mutant rice high in resistant starch,Journal of Agricultural&Food Chemistry,54(2):523-528
Yang R.,Sun C.,Bai J.,Luo Z.,Shi B.,Zhang J.,Yan W.,and Piao Z.,2012,A putative gene sbe3-rs for resistant starch mutated from SBE3 for starch branching enzyme in rice(O-ryza sativa L.),PLoS One,7(8):e43026
Zhou H.,Wang L.,Liu G.,Meng X.,Jing Y.,Shu X.,Kong X.,Sun J.,Yu H.,and Smith S.M.,2016,From the cover:critical roles of soluble starch synthase SSⅢa and granule-bound starch synthase Waxy in synthesizing resistant starch in rice,Proc.Natl.Acad.Sci.USA,113(45):12844-12849
Zhu H.M.,Bai J.J.,Wang H.,Li M.B.,Hao Z.S.,and Pu Z.Z.,2010,Starch properties of japonica rice enriched with resistant starch,Zhonggu o Nongxue Tonbao(Chinese Agricultural Science Bulletin),26(14):108-112(朱辉明,白建江,王慧,李茂柏,郝再彬,朴钟泽,2010,高抗性淀粉粳稻新品系稻米淀粉特性,中国农学通报,26(14):108-112)
Ai Y.,Hasjim J.,and Jane J.L.,2013,Effects of lipids on enzymatic hydrolysis and physical properties of starch,Carbohydrate Polymers,92(1):120-127
Baghurst P.A.,Baghurst K.I.,and Record S.J.,1996,Dietary fibre,non-starch polysaccharides and resistant starch:a review,Food Australia,48(3):S3-S35
Ball S.,Guan H.P.,James M.,Myers A.,Keeling P.,Mouille G.Buléon A.,Colonna P.,and Preiss J.,1996,From glycogen to amylopectin:a model for the biogenesis of the plant starch granule,Cell,86(3):349-352
Bao J.S.,Zhou X.,Xu F.F.,He Q.,and Park Y.J.,2017,Genomewide association study of the resistant starch content in rice grains,Starch/St覿rke,69(7-8):1600343
Bird A.R.,Brown I.L.,and Topping D.L.,2000,Starches,resistant starches,the gut microflora and human health,Curr.Issues in Intest.Microbiol.,1(1):25-37
Brouns F.,Kettlitz B.,and Arrigoni E.,2002,Resistant starch and"the butyrate revolution",Trends in Food Science&Technology,13(8):251-261
Chen L.,Magliano D.J.,and Zimmet P.Z.,2011,The worldwide epidemiology of type 2 diabetes mellitus-present and future perspectives,Nat.Rev.Endocrinol.,8(4):228-236
Dupuis J.H.,Liu Q.,and Yada R.Y.,2015,Methodologies for increasing the resistant starch content of food starches:a review,Comprehensive Reviews in Food Science&Food Safety,13(6):1219-1234
Englyst H.,Wiggins H.S.,and Cummings J.H.,1982,Determination of the non-starch polysaccharides in plant foods by gas-liquid chromatography of constituent sugars as alditol acetates,Analyst,107(1272):307-318
Fitzgerald M.A.,Rahman S.,Resurreccion A.P.,Concepcion J.,Daygon V.D.,Dipti S.S.,Kabir K.A.,Klingner B.,Morell M.K.,and Bird A.R.,2011,Identification of a major genetic determinant of glycaemic index in rice,Rice,4(2):66-74
Guo C.F.,Li J.Y.,and Zhang S.W.,2006,The research development on the physiological function of resistant starch,Shipin Keji(Food Science and Technology),31(2):1-3(郭春锋,李婧妍,张守文,2006,抗性淀粉生理功能研究进展,食品科技,31(2):1-3)
Han J.A.,and Bemiller J.N.,2007,Preparation and physical characteristics of slowly digesting modified food starches,Carbohydrate Polymers,67(3):366-374
Hannah L.C.,and James M.,2008,The complexities of starch biosynthesis in cereal endosperms,Curr.Opin.Biotechnol.,19(2):160-165
Hennen-Bierwagen T.A.,Lin Q.,Grimaud F.,Planchot V.,Keeling P.L.,James M.G.,and Myers A.M.,2009,Proteins from multiple metabolic pathways associate with starch biosynthetic enzymes in high molecular weight complexes:a model for regulation of carbon allocation in maize amyloplasts,Plant Physiol.,149(3):1541-1559
Hu P.,Zhao H.J.,Duan Z.Y.,Zhang L.L.,and Wu D.X.,2004,Starch digestibility and the estimated glycemic score of different types of rice differing in amylose contents,Journal of Cereal Science,40(3):231-237
James M.G.,Denyer K.,and Myers A.M.,2003,Starch synthesis in the cereal endosperm,Curr.Opin.Plant Biol.,6(3):215-222
Kang H.G.,Park S.H.,Matsuoka M.,and An G.H.,2005White-core endosperm floury endosperm-4 in rice is generated by knockout mutations in the C4-type pyruvate orthophosphate dikinase gene(OsPPDKB),Plant J.,42(6):901-911
Luo X.,Huang J.F.,Zhu Y.S.,Xie H.G.,Wu F.X.,Zhang M.Q.Zhang J.F.,and Xie H.A.,2014,Genetic analysis of high resistant starch characteristics for rice variety gongmi 3(Oryza sativa ssp.indica),Nongye Shengwu Jishu Xuebao(Journa of Agricultural Biotechnology),22(1):10-16(罗曦,黄锦峰朱永生,谢鸿光,吴方喜,张木清,张建福,谢华安,2014水稻功米3号高抗性淀粉性状的遗传分析,农业生物技术学报,22(1):10-16)
Myers A.M.,Morell M.K.,James M.G.,and Ball S.G.,2000,Recent progress toward understanding biosynthesis of the amylopectin crystal,Plant Physiol.,122(4):989-997
Nakamura Y.,2002,Towards a better understanding of the metabolic system for amylopectin biosynthesis in plants:rice endosperm as a model tissue,Plant&Cell Physiology,43(7):718-725
Ordonio R.L.,and Matsuoka M.,2016,Increasing resistant starch content in rice for better consumer health,Proc.Natl.Acad Sci.USA,113(45):12616-12618
Raben A.,Tagliabue A.,Christensen N.J.,Madsen J.,Holst J.J.and Astrup A.,1994,Resistant starch:the effect on postprandial glycemia,hormonal response,and satiety,American Journal of Clinical Nutrition,60(4):544-551
Raigond P.,Ezekiel R.,and Raigond B.,2015,Resistant starch in food:a review,J.Sci.Food Agric.,95(10):1968-1978
Sun Z.M.,Bai J.J.,Zhu H.M.,Sun C.L.,Pu Z.Z.,and Hao Z.S.2012,Effects of mixing ratio and processing methods on resistant starch content of products of rice'Jiangtangdao 1'Henongxue Bao(Journal of Nuclear Agricultural Sciences)26(2):318-323(孙志敏,白建江,朱辉明,孙春龙,朴钟泽郝再彬,2012,混配比例及加工方式对“降糖稻1号”稻米产品抗性淀粉含量的影响,核农学报,26(2):318-323)
Tian Z.X.,Qian Q.,Liu Q.Q.,Yan M.X.,Liu X.F.,Yan C.J.,Liu G.F.,Gao Z.Y.,Tang S.Z.,and Zeng D.L.,2009,Allelic diversities in rice starch biosynthesis lead to a diverse array of rice eating and cooking qualities,Proc.Natl.Acad.Sci.US-A,106(51):21760-21765
Tran N.A.,Daygon V.D.,Resurreccion A.P.,Cuevas R.P.,Corpuz H.M.,and Fitzgerald M.A.,2011,A single nucleotide polymorphism in the Waxy gene explains a significant component of gel consistency,Theor.Appl.Genet.,123(4)519-525
Van de Wal M.,D'Hulst C.,Vincken J.P.,Buléon A.,Visser R.,and Ball S.,1998,Amylose is synthesized in vitro by extension of and cleavage from amylopectin,J.Biol.Chem.,273(35):22232-22240
Witt T.,Gidley M.J.,and Gilbert R.G.,2010,Starch digestion mechanistic information from the time evolution of molecular size distributions,J.Agric.Food Chem.,58(14):8444-8452
Wu W.,Liu X.,Yang C.Z.,Shu X.L.,and Wu D.X.,2006,Progresses in the study of resistant starch and functional rice for prevention of diabetes and hyperlipidemia,Henongxue Bao(Journal of Nuclear Agricultural Sciences),20(1):60-63(吴伟,刘鑫,杨朝柱,舒小丽,吴殿星,2006,抗性淀粉及预防糖尿病和肥胖症功能稻米研究进展,核农学报,20(1):60-63)
Yamada Y.,Hosoya S.,Nishimura S.,Tanaka T.,Kajimoto Y.,Nishimura A.,and Kajimoto O.,2005,Effect of bread containing resistant starch on postprandial blood glucose levels in humans,Biosci.Biotechnol.Biochem.,69(3):559-566
Yang C.Z.,Li C.S.,Shu X.L.,Zhang Z.Z.,Zhang L.,Zhao H.J.,Ma C.X.,and Wu D.X.,2005,Starch properties of rice mutant enriched with resistant starch,Zhongguo Shuidao Kexue(Chinese Journal of Rice Science),19(6):516-520(杨朝柱,李春寿,舒小丽,张志转,张磊,赵海军,马传喜,吴殿星,2005,富含抗性淀粉水稻突变体的淀粉特性,中国水稻科学,19(6):516-520)
Yang C.Z.,Shu X.L.,Zhang L.L.,Wang X.Y.,Zhao H.J.,Ma C.X.,and Wu D.X.,2006,Starch properties of mutant rice high in resistant starch,Journal of Agricultural&Food Chemistry,54(2):523-528
Yang R.,Sun C.,Bai J.,Luo Z.,Shi B.,Zhang J.,Yan W.,and Piao Z.,2012,A putative gene sbe3-rs for resistant starch mutated from SBE3 for starch branching enzyme in rice(O-ryza sativa L.),PLoS One,7(8):e43026
Zhou H.,Wang L.,Liu G.,Meng X.,Jing Y.,Shu X.,Kong X.,Sun J.,Yu H.,and Smith S.M.,2016,From the cover:critical roles of soluble starch synthase SSⅢa and granule-bound starch synthase Waxy in synthesizing resistant starch in rice,Proc.Natl.Acad.Sci.USA,113(45):12844-12849
Zhu H.M.,Bai J.J.,Wang H.,Li M.B.,Hao Z.S.,and Pu Z.Z.,2010,Starch properties of japonica rice enriched with resistant starch,Zhonggu o Nongxue Tonbao(Chinese Agricultural Science Bulletin),26(14):108-112(朱辉明,白建江,王慧,李茂柏,郝再彬,朴钟泽,2010,高抗性淀粉粳稻新品系稻米淀粉特性,中国农学通报,26(14):108-112)