氮肥对水稻胚乳淀粉品质、相关酶活性及基因表达量的影响
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摘要
【目的】旨在为阐明灌浆成熟期氮素营养对水稻淀粉品质影响机理以及建立优质高产水稻栽培技术提供理论依据。【方法】选用4个籽粒直链淀粉含量差异显著的粳稻品种,通过盆栽试验研究氮素营养对稻米淀粉组分和蛋白质含量及稻米蛋白质水解对淀粉黏滞特性的影响,并分析氮素营养对灌浆过程中籽粒蔗糖合酶(SuSy)、蔗糖磷酸合酶(SPS)、蔗糖酸性转化酶(AI)活性及OsGBSSⅠ、Os ISAⅠ、OsSBEⅠ、OsSBEⅡ基因转录表达量的影响。【结果】结果表明,籽粒支链淀粉含量对氮素营养很敏感,灌浆成熟期氮素营养能改变籽粒淀粉组分含量;施氮条件下,高直链淀粉含量品种蒸煮食味品质下降更加明显,去除稻米蛋白质可明显提高稻米黏滞特性,蛋白质对淀粉黏滞特性的影响很大;增加灌浆成熟期氮素营养能显著或极显著提高籽粒Su Sy和AI活性,显著抑制籽粒SPS活性;灌浆成熟期氮素营养能改变灌浆不同时期籽粒OsGBSSⅠ、OsISAⅠ和OsSBEⅡ基因转录表达量,以致灌浆过程中这些基因的转录表达量变化动态发生改变,但OsSBEⅠ基因转录表达量不因氮素营养而发生改变;受氮素营养的影响,灌浆起始期籽粒OsGBSSⅠ基因表达量明显上调,而灌浆中后期明显下调;氮素营养明显抑制灌浆成熟期籽粒OsI SAⅠ基因和灌浆中后期的OsS BEⅠ基因转录表达,显著提高灌浆前期和中期的籽粒OsSBEⅠ基因转录表达量;氮素营养能抑制灌浆起始期籽粒OsSBEⅡ基因的转录表达,而提高灌浆中后期的基因转录表达。【结论】灌浆成熟期氮素营养除了通过蛋白质含量对淀粉品质产生影响外,还通过调控淀粉合成相关的酶活性和基因表达量等生理环节对淀粉含量和精细结构起作用,最终改变稻米黏滞特性和食味品质。
【Objective】The research aims to lay a theoretical basis for clarifying the influencing mechanism of nitrogen fertilization during grain filling period on rice starch quality.【Method】Four japonica rice varieties with significant difference in amylose content in grains were selected. Pot experiments were conducted to study the effect of nitrogen application on the starch component and protein content in rice grains, and the protein hydrolysis of rice grains on starch viscosity properties, and to analyze the effect of nitrogen nutrition on the activity of sucrose synthase(SuSy), sucrose phosphate synthase(SPS) and acid invertase(AI) and the m RNA expression levels of OsGBSSⅠ, Os ISAⅠ, OsSBEⅠand OsSBEⅡ genes of rice grains during grain filling period.【Results】The results showed that the grain amylopectin content was sensitive to nitrogen nutrition, and nitrogen nutrition during grain filling period changed the content of starch components in grains. The cooking and eating quality of high amylose content varieties degraded even more. The removal of rice protein could significantly improve the viscosity of rice, and the starch composition had a great influence on the viscosity properties of rice starch. Increasing nitrogen nutrition at grain filling stage significantly or highly significantly increased the activity of Su Sy and AI in grains, and significantly inhibits the activity of SPS. The nitrogen nutrition at grain filling stage changed the m RNA expression level of Os GBSSⅠ, Os ISAⅠ and OsSBEⅡ of rice grains at different stages of grain filling, so that the change dynamics of the m RNA expression quantity of these genes changed during grain filling period except OsSBEⅠ. Due to the influence of nitrogen nutrition, at the beginning of grain filling, Os GBSSⅠ expression level was upregulated significantly, but downregulated significantly in the middle and late stages of grain filling. Nitrogen fertilization could significantly inhibit the transcription expression quantity of Os ISAⅠ of grains at grain filling stage and OsSBEⅠ of grains in the middle and late stages of grain filling, and could significantly improve the m RNA expression quantity of OsSBEⅠ of grains in the early and middle stages of grain filling. Nitrogen nutrition could inhibit the transcription expression quantity of OsSBEⅡ gene of grains at the beginning of grain filling, and improve the transcription expression quantity of genes in the middle and late stages of grain filling.【Conclusion】In addition to the effect of protein content on starch quality during grain filling and ripening, nitrogen nutrition also affected starch content and fine structure by regulating enzyme activities and gene expression related to starch synthesis. Ultimately, it changed the rice viscosity and taste quality.
【Objective】The research aims to lay a theoretical basis for clarifying the influencing mechanism of nitrogen fertilization during grain filling period on rice starch quality.【Method】Four japonica rice varieties with significant difference in amylose content in grains were selected. Pot experiments were conducted to study the effect of nitrogen application on the starch component and protein content in rice grains, and the protein hydrolysis of rice grains on starch viscosity properties, and to analyze the effect of nitrogen nutrition on the activity of sucrose synthase(SuSy), sucrose phosphate synthase(SPS) and acid invertase(AI) and the m RNA expression levels of OsGBSSⅠ, Os ISAⅠ, OsSBEⅠand OsSBEⅡ genes of rice grains during grain filling period.【Results】The results showed that the grain amylopectin content was sensitive to nitrogen nutrition, and nitrogen nutrition during grain filling period changed the content of starch components in grains. The cooking and eating quality of high amylose content varieties degraded even more. The removal of rice protein could significantly improve the viscosity of rice, and the starch composition had a great influence on the viscosity properties of rice starch. Increasing nitrogen nutrition at grain filling stage significantly or highly significantly increased the activity of Su Sy and AI in grains, and significantly inhibits the activity of SPS. The nitrogen nutrition at grain filling stage changed the m RNA expression level of Os GBSSⅠ, Os ISAⅠ and OsSBEⅡ of rice grains at different stages of grain filling, so that the change dynamics of the m RNA expression quantity of these genes changed during grain filling period except OsSBEⅠ. Due to the influence of nitrogen nutrition, at the beginning of grain filling, Os GBSSⅠ expression level was upregulated significantly, but downregulated significantly in the middle and late stages of grain filling. Nitrogen fertilization could significantly inhibit the transcription expression quantity of Os ISAⅠ of grains at grain filling stage and OsSBEⅠ of grains in the middle and late stages of grain filling, and could significantly improve the m RNA expression quantity of OsSBEⅠ of grains in the early and middle stages of grain filling. Nitrogen nutrition could inhibit the transcription expression quantity of OsSBEⅡ gene of grains at the beginning of grain filling, and improve the transcription expression quantity of genes in the middle and late stages of grain filling.【Conclusion】In addition to the effect of protein content on starch quality during grain filling and ripening, nitrogen nutrition also affected starch content and fine structure by regulating enzyme activities and gene expression related to starch synthesis. Ultimately, it changed the rice viscosity and taste quality.
引文
[1]沈鹏,金正勋,罗秋香,金学泳,孙艳丽.水稻灌浆过程中籽粒淀粉合成关键酶活性与蒸煮食味品质的关系.中国水稻科学,2006,20(1):58-64.Shen P,Jin Z X,Luo Q X,Jin X Y,Sun Y L.Relationship between activity of key starch synthetic enzymes during grain filling and quality of eating and cooking in rice.Chin J Rice Sci,2006,20(1):58-64.(in Chinese with English abstract)
[2]李晓光,金正勋,刘海英,赵书宇,徐美兰,张忠臣,沈鹏,张海彬.水稻杂种后代籽粒直链淀粉和蛋白质含量选择对产量和品质性状的影响.东北农业大学学报,2011,42(4):13-17.Li X G,Jin Z X,Liu H Y,Zhao S Y,Xu M L,Zhang Z C,Shen P,Zhang H B.Effect of amylose and protein content selection on yield and quality traits of hybrid progenies in rice.J Northeast Agric Univ,2011,42(4):13-17.(in Chinese with English abstract)
[3]张欣,施利利,刘晓宇,丁得亮,王松文,崔晶.不同施肥处理对水稻产量、食味品质及蛋白质组分的响.中国农学通报,2010,26(4):104-108.Zhang X,Shi L L,Liu X Y,Ding D L,Wang S W,Cui J.Effect of different fertilizer treatments on rice yield,grain quality and protein fraction content.Chin Agric Sci Bull,2010,26(4):104-108.(in Chinese with English abstract)
[4]谢黎虹,罗炬,唐绍清,陈能,焦桂爱,邵高能,魏祥进,胡培松.蛋白质影响水稻米饭食味品质的机理.中国水稻科学,2013,27(1):91-96.Xie L H,Luo J,Tang S Q,Chen N,Jiao G A,Shao G N,Wei X J,Hu P S.Proteins affect rice eating quality properties and its mechanism.Chin J Rice Sci,2013,27(1):91-96.(in Chinese with English abstract)
[5]阳剑,时亚文,李宙炜,陶优生,唐启源.水稻碳氮代谢研究进展.作物研究,2011,25(4):383-387.Yang J,Shi Y W,Li Z Y,Tao Y S,Tang Q Y.Research progress of rice carbon and nitrogen metabolism.Crop Res,2011,25(4):383-387.(in Chinese with English abstract)
[6]朱方旭,郭雪冬,同拉嘎,张玉磊,潘冬,李明月,李丹,张忠臣,金正勋.蘖穗氮肥追施比例对水稻灌浆成熟期Rubisco和GS同工型基因表达量的影响.植物营养与肥料学报,2017,23(2):324-332.Zhu F X,Guo X D,Tong L G,Zhang Y L,Pan D,Li M Y,Li D,Zhang Z C,Jin Z X.Expression response of Rubisco and GS isoform gene to the ratio of tillering and heading nitrogen fertilization at rice filling stage.J Plant Nutr Fert,2017,23(2):324-332.(in Chinese with English abstract)
[7]金正勋,郭雪冬,朱方旭,张忠臣.灌浆成熟期氮素营养对粳稻品质性状及籽粒蔗糖代谢相关酶活性影响.东北农业大学学报,2016,47(9):1-6.Jin Z X,Guo X D,Zhu F X,Zhang Z C.Effect of nitrogen nutrition in japonica quality characters and grain sucrose-metabolizing enzymes activity during grouting mature stage.J Northeast Agric Univ,2016,47(9):1-6.(in Chinese with English abstract)
[8]从夕汉,施伏芝,阮新民,罗玉祥,马廷臣,罗志祥.氮肥水平对不同基因型水稻氮素利用率、产量和品质的影响.应用生态学报,2017,28(4):1219-1226.Cong X H,Shi F Z,Ruan X M,Luo Y X,Ma T C,Luo ZX.Effects of nitrogen fertilizer application rate on nitrogen use efficiency and grain yield and quality of different rice varieties.Chin J Appl Ecol,2017,28(4):1219-1226.(in Chinese with English abstract)
[9]王龙,仲维君,张丽微,黄成亮,潘世驹,姜玉伟,赵婷婷,宋泽,周健,钱永德.结实期施氮对水稻空育131品质的影响.四川农业大学学报,2016,31(1):19-23.Wang L,Zhong W J,Zhang L W,Huang C L,Pan S J,Jiang Y W,Zhao T T,Song Z,Zhou J,Qian Y D.Effect of nitrogen fertilizer on the quality of the Kongyu 131 in seed setting stage.J Sichuan Agric Univ,2016,31(1):19-23.(in Chinese with English abstract)
[10]李运祥,王忠,顾蕴洁,陈召洲.施氮处理对稻米淀粉积累的影响.南京师大学报:自然科学版,2003,26(3):31-35.Li Y X,Wang Z,Gu Y J,Chen Z Z.Effects of various nitrogen treatments on starch accumulation in caryopsis during the rice grain filling.J Nanjing Nor Univ:Nat Sci Ed,2003,26(3):31-35.(in Chinese with English abstract)
[11]朱立楠,刘海英,孙璐璐,孙涛,郭雪冬,朱方旭,张忠臣,金正勋.水稻灌浆过程中胚乳异淀粉酶基因家族表达特性及其与淀粉含量间关系分析.中国水稻科学,2015,29(5):528-534.Zhu L N,Liu H Y,Sun L L,Sun T,Guo X D,Zhu F X,Zhang Z C,Jin Z X.Analysis of expression characteristics of isoamylase and the correlation with starch content during grain filling in rice.Chin J Rice Sci,2015,29(5):528-534.(in Chinese with English abstract)
[12]杜晓东,赵宏伟,王敬国,刘化龙,杨亮,许晶,宋谨同.氮素运筹对寒地粳稻淀粉合成关键酶活性及淀粉积累的影响.作物学报,2012,38(1):159-167.Du X D,Zhao H W,Wang J G,Liu H L,Yang L,Xu J,Song J T.Changes in starch accumulation and activity of enzymes associated with starch synthesis under different nitrogen applications in japonica rice in cold region.Acta Agron Sin,2012,38(1):159-167.(in Chinese with English abstract)
[13]中华人民共和国农业部.中华人民共和国农业部部颁标准NY147-88.米质测定方法.北京:中国标准出版社,1988.Ministry of Agriculture,the Peoples Republic of China.Measuring Method of Rice Quality Characteristics.NY147-88.Beijing:China Standards Press,1988.(in Chinese)
[14]何照范.谷物淀粉组份分离及测试方法评述.粮食储藏,1985,65(6):32-38.He Z F,A review of the separation and test methods of cereal starch.Grain Stor,1985,65(6):32-38.(in Chinese)
[15]American Association of Cereal Chemist(AACC).Approved methods of analysis.11th edn.Methods 61-02.01.Determination of the Pasting Properties of Rice with the Rapid Visco Analyser.The Association:St.Paul,MN,1999.
[16]杨玉玲,许时婴.籼米为基质的脂肪替代品-糊化工艺研究.中国粮油学报,2002,17(6):48-50.Yang Y L,Xu S Y.Rice based fat substitute:Study on gelatinization.Chin Cereals Oils Assoc,2002,17(6):48-50.(in Chinese with English abstract)
[17]Chrastil J.Protein-starch interactions in rice grains.Influence of storage on oryzenin and starch.J Agric Food Chem,1990,38(9):1804-1809.
[18]金正勋,朱方旭,郭雪冬,张忠臣.不同施氮方法对粳稻灌浆成熟期蔗糖代谢相关酶活性及品质性状影响.东北农业大学学报,2016,47(6):1-7.Jin Z X,Zhu F X,Guo X D,Zhang Z C.Effect of different nitrogen application methods on metabolism correlation enzymes and quality in rice.J Northeast Agric Univ,2016,47(6):1-7.(in Chinese with English abstract)
[19]Rajeevan M S,Ranamukhaarachchi D G,Vernon S D,Unger E R.Use of real-time quantitative PCR to validate the results of c DNA array and differential display PCRtechnologies.Methods(San Diego,Calif.),2001,25(4):443-451.
[20]孙永健,孙园园,李旭毅,郭翔,马均.水氮互作下水稻氮代谢关键酶活与氮素利用的关系.作物学报,2009,35(11):2055-2063Sun Y J,Sun Y Y,Li X Y,Guo X,Ma J.Relationship of activities of key enzymes involved in nitrogen metabolism with nitrogen utilization in rice under water-nitrogen interaction.Acta Agron Sin,2009,35(11):2055-2063.(in Chinese with English abstract)
[21]苗红霞,孙佩光,张凯星,金志强,徐碧玉.植物颗粒结合淀粉合成酶(GBSS)基因的表达调控机制研究进展.生物技术通报,2016,32(3):18-23.Miao H X,Sun P G,Zhang K X,Jin Z Q,Xu B Y.Research progress on expression regulation mechanism of genes encoding granule-bound starch synthase in plants.Biotech Bull,2016,32(3):18-23.(in Chinese with English abstract)
[22]陈书强,杨丽敏,赵海新,杜晓东,周通,薛菁芳,单莉莉,王翠,张献国.穗粒肥用氮比例对寒地不同类型水稻产量和品质的影响.中国稻米.2017,23(4):151-156.Chen S Q,Yang L M,Zhao H X,Du X D,Zhou T,Xue JF,Shan L L,Wang C,Zhang X G.Effects of the spike-grain nitrogen ratio on yield and quality of different types of rice in cold region.China Rice,2017,23(4):151-156.(in Chinese)
[23]Kim W,Johnson J W,Graybosch R A,Gaines C S.Physicochemical properties and end-use quality of wheat starch as a function of waxy protein alleles.J Cereal Sci,2003,37(2):195-204.
[24]言普,沈文涛,黎小瑛,周鹏.植物hp RNA干扰载体构建的研究进展.生物技术通报,2013(9):7-12.Yan P,Shen W T,Li X Y,Zhou P.Progress in Construction of hp RNA vector for plant RNAi.Biotechnol Bull,2013(9):7-12.(in Chinese with English abstract)
[25]Sasaki T,Yasui T,Matsuki J.Effect of amylose content on gelatinization,retrogradation,and pasting properties of starches from waxy and nonwaxy wheat and their F1seeds.Cereal Chem,2000,77(1):58-63.
[26]Wang J C,Xu H,Zhu Y,Liu Q Q,Cai X L.Osb ZIP58,a basic leucine zipper transcription factor,regulates starch biosynthesis in rice endosperm//2012 National Conference on Plant Biology.2012:3453-3466.
[27]方先文,姜东,戴廷波,曹卫星.小麦淀粉组分的积累规律.江苏农业学报,2002,18(3):139-142.Fang X W,Jiang D,Dai T B,Cao W X.Accumulation regularity of starch components in wheat varieties.Jiangsu J Agric Sci,2002,18(3):139-142.(in Chinese with English abstract)
[28]Nishi A,Nakamura Y,Tanaka N,Satoh H.Organ Specificity of isoforms of starch branching enzyme(Q-enzyme)in rice.Plant Cell Physiol,1992,33(7):985-991.
[29]Satoh H,Nishi A,Yamashita K,Takemoto Y.Alteration of the structural properties of starch components by the lack of an isoforms of starch branching enzyme in rice seeds.J Biol Chem,1993,268(25):19084-19091.
[30]Kawasaki T,Mizuno K,Baba T,Shimada H.Molecular analysis of the gene encoding a rice starch branching enzyme.Mol Gen Genet,1993,237(1):10-16.
[31]Guan H P,Preiss J.Differentiation of the properties of the branching isozymes from maize(Zea mays).Plant Physiol,1993,102(4):1269-1273.
[32]Morell M K,Blennow A,Kosar-Hasherni B,Samuel MS.Differential expression and Properties of starch branching enzyme isoforms in developing wheat endosperm.Plant Physiol,1997,113(1):201-208.
[33]Satoh H,Nishi A,Yamashita K,Takemoto Y,Tanaka Y,Hosaka Y,Sakurai A,Fujita N,Nakamura Y.Starch-branching enzyme I deficient mutation specifically affects the structure and properties of starch in rice endosperm.Plant Physi,2003,133:1111-1121.
[34]Utsumi Y,Utsumi C,Sawada T,Fujita N,Nakamura Y.Functional diversity of isoamylase oligomers:The ISA1homo-oligomer is essential for amylopectin biosynthesis in rice endosperm.Plant Physiol,2011,156(1):61-77.
[35]Kubo A,Rahman S,Utsumi Y.Complementation of sugary-1phenotype in rice endosperm with the wheat isoamy-lase1gene supports a direct role forisoamylase1 in amylopectin biosynthesis.Plant Physiol,2005,137(1):43-56.
[36]钟连进,董虎,蔡小波,封言柠,任凭,程方民.控制水稻胚乳淀粉合成代谢若干关键酶基因对花后高温的响应表达.应用生态学报,2012,23(3):745-750.Zhong L J,Dong H,Cai X B,Feng Y N,Ren P,Cheng FM.Gene expression of the key enzymes controlling starch synthesis and metabolism in rice grain endosperm under effects of high temperature after anthesis.Chin JAppl Ecol,2012,23(3):745-750.(in Chinese with English abstract)
[37]钟连进,程方民,张国平,孙宗修.灌浆结实期不同温度下早籼稻米淀粉链长分布与结构特征差异分析.中国农业科学,2005,38(2):272-276.Zhong L J,Cheng F M,Zhang G P,Sun Z X.Differences in starch chain length distribution and structure characteristics of early-indica rice under different temperature treatments during grain-filling.Sci Agric Sin,2005,38(2):272-276.(in Chinese with English abstract)
[38]Delatte T,Trevisan M,Parker M L,Zeeman S C.Arabidopsis mutants Atisa1 and Atisa2 have identical phenotypes and lack the same multimeric isoamylase,which influences the branch point distribution of amylopectin during starch synthesis.Plant J,2005,41(6):815-830.
[2]李晓光,金正勋,刘海英,赵书宇,徐美兰,张忠臣,沈鹏,张海彬.水稻杂种后代籽粒直链淀粉和蛋白质含量选择对产量和品质性状的影响.东北农业大学学报,2011,42(4):13-17.Li X G,Jin Z X,Liu H Y,Zhao S Y,Xu M L,Zhang Z C,Shen P,Zhang H B.Effect of amylose and protein content selection on yield and quality traits of hybrid progenies in rice.J Northeast Agric Univ,2011,42(4):13-17.(in Chinese with English abstract)
[3]张欣,施利利,刘晓宇,丁得亮,王松文,崔晶.不同施肥处理对水稻产量、食味品质及蛋白质组分的响.中国农学通报,2010,26(4):104-108.Zhang X,Shi L L,Liu X Y,Ding D L,Wang S W,Cui J.Effect of different fertilizer treatments on rice yield,grain quality and protein fraction content.Chin Agric Sci Bull,2010,26(4):104-108.(in Chinese with English abstract)
[4]谢黎虹,罗炬,唐绍清,陈能,焦桂爱,邵高能,魏祥进,胡培松.蛋白质影响水稻米饭食味品质的机理.中国水稻科学,2013,27(1):91-96.Xie L H,Luo J,Tang S Q,Chen N,Jiao G A,Shao G N,Wei X J,Hu P S.Proteins affect rice eating quality properties and its mechanism.Chin J Rice Sci,2013,27(1):91-96.(in Chinese with English abstract)
[5]阳剑,时亚文,李宙炜,陶优生,唐启源.水稻碳氮代谢研究进展.作物研究,2011,25(4):383-387.Yang J,Shi Y W,Li Z Y,Tao Y S,Tang Q Y.Research progress of rice carbon and nitrogen metabolism.Crop Res,2011,25(4):383-387.(in Chinese with English abstract)
[6]朱方旭,郭雪冬,同拉嘎,张玉磊,潘冬,李明月,李丹,张忠臣,金正勋.蘖穗氮肥追施比例对水稻灌浆成熟期Rubisco和GS同工型基因表达量的影响.植物营养与肥料学报,2017,23(2):324-332.Zhu F X,Guo X D,Tong L G,Zhang Y L,Pan D,Li M Y,Li D,Zhang Z C,Jin Z X.Expression response of Rubisco and GS isoform gene to the ratio of tillering and heading nitrogen fertilization at rice filling stage.J Plant Nutr Fert,2017,23(2):324-332.(in Chinese with English abstract)
[7]金正勋,郭雪冬,朱方旭,张忠臣.灌浆成熟期氮素营养对粳稻品质性状及籽粒蔗糖代谢相关酶活性影响.东北农业大学学报,2016,47(9):1-6.Jin Z X,Guo X D,Zhu F X,Zhang Z C.Effect of nitrogen nutrition in japonica quality characters and grain sucrose-metabolizing enzymes activity during grouting mature stage.J Northeast Agric Univ,2016,47(9):1-6.(in Chinese with English abstract)
[8]从夕汉,施伏芝,阮新民,罗玉祥,马廷臣,罗志祥.氮肥水平对不同基因型水稻氮素利用率、产量和品质的影响.应用生态学报,2017,28(4):1219-1226.Cong X H,Shi F Z,Ruan X M,Luo Y X,Ma T C,Luo ZX.Effects of nitrogen fertilizer application rate on nitrogen use efficiency and grain yield and quality of different rice varieties.Chin J Appl Ecol,2017,28(4):1219-1226.(in Chinese with English abstract)
[9]王龙,仲维君,张丽微,黄成亮,潘世驹,姜玉伟,赵婷婷,宋泽,周健,钱永德.结实期施氮对水稻空育131品质的影响.四川农业大学学报,2016,31(1):19-23.Wang L,Zhong W J,Zhang L W,Huang C L,Pan S J,Jiang Y W,Zhao T T,Song Z,Zhou J,Qian Y D.Effect of nitrogen fertilizer on the quality of the Kongyu 131 in seed setting stage.J Sichuan Agric Univ,2016,31(1):19-23.(in Chinese with English abstract)
[10]李运祥,王忠,顾蕴洁,陈召洲.施氮处理对稻米淀粉积累的影响.南京师大学报:自然科学版,2003,26(3):31-35.Li Y X,Wang Z,Gu Y J,Chen Z Z.Effects of various nitrogen treatments on starch accumulation in caryopsis during the rice grain filling.J Nanjing Nor Univ:Nat Sci Ed,2003,26(3):31-35.(in Chinese with English abstract)
[11]朱立楠,刘海英,孙璐璐,孙涛,郭雪冬,朱方旭,张忠臣,金正勋.水稻灌浆过程中胚乳异淀粉酶基因家族表达特性及其与淀粉含量间关系分析.中国水稻科学,2015,29(5):528-534.Zhu L N,Liu H Y,Sun L L,Sun T,Guo X D,Zhu F X,Zhang Z C,Jin Z X.Analysis of expression characteristics of isoamylase and the correlation with starch content during grain filling in rice.Chin J Rice Sci,2015,29(5):528-534.(in Chinese with English abstract)
[12]杜晓东,赵宏伟,王敬国,刘化龙,杨亮,许晶,宋谨同.氮素运筹对寒地粳稻淀粉合成关键酶活性及淀粉积累的影响.作物学报,2012,38(1):159-167.Du X D,Zhao H W,Wang J G,Liu H L,Yang L,Xu J,Song J T.Changes in starch accumulation and activity of enzymes associated with starch synthesis under different nitrogen applications in japonica rice in cold region.Acta Agron Sin,2012,38(1):159-167.(in Chinese with English abstract)
[13]中华人民共和国农业部.中华人民共和国农业部部颁标准NY147-88.米质测定方法.北京:中国标准出版社,1988.Ministry of Agriculture,the Peoples Republic of China.Measuring Method of Rice Quality Characteristics.NY147-88.Beijing:China Standards Press,1988.(in Chinese)
[14]何照范.谷物淀粉组份分离及测试方法评述.粮食储藏,1985,65(6):32-38.He Z F,A review of the separation and test methods of cereal starch.Grain Stor,1985,65(6):32-38.(in Chinese)
[15]American Association of Cereal Chemist(AACC).Approved methods of analysis.11th edn.Methods 61-02.01.Determination of the Pasting Properties of Rice with the Rapid Visco Analyser.The Association:St.Paul,MN,1999.
[16]杨玉玲,许时婴.籼米为基质的脂肪替代品-糊化工艺研究.中国粮油学报,2002,17(6):48-50.Yang Y L,Xu S Y.Rice based fat substitute:Study on gelatinization.Chin Cereals Oils Assoc,2002,17(6):48-50.(in Chinese with English abstract)
[17]Chrastil J.Protein-starch interactions in rice grains.Influence of storage on oryzenin and starch.J Agric Food Chem,1990,38(9):1804-1809.
[18]金正勋,朱方旭,郭雪冬,张忠臣.不同施氮方法对粳稻灌浆成熟期蔗糖代谢相关酶活性及品质性状影响.东北农业大学学报,2016,47(6):1-7.Jin Z X,Zhu F X,Guo X D,Zhang Z C.Effect of different nitrogen application methods on metabolism correlation enzymes and quality in rice.J Northeast Agric Univ,2016,47(6):1-7.(in Chinese with English abstract)
[19]Rajeevan M S,Ranamukhaarachchi D G,Vernon S D,Unger E R.Use of real-time quantitative PCR to validate the results of c DNA array and differential display PCRtechnologies.Methods(San Diego,Calif.),2001,25(4):443-451.
[20]孙永健,孙园园,李旭毅,郭翔,马均.水氮互作下水稻氮代谢关键酶活与氮素利用的关系.作物学报,2009,35(11):2055-2063Sun Y J,Sun Y Y,Li X Y,Guo X,Ma J.Relationship of activities of key enzymes involved in nitrogen metabolism with nitrogen utilization in rice under water-nitrogen interaction.Acta Agron Sin,2009,35(11):2055-2063.(in Chinese with English abstract)
[21]苗红霞,孙佩光,张凯星,金志强,徐碧玉.植物颗粒结合淀粉合成酶(GBSS)基因的表达调控机制研究进展.生物技术通报,2016,32(3):18-23.Miao H X,Sun P G,Zhang K X,Jin Z Q,Xu B Y.Research progress on expression regulation mechanism of genes encoding granule-bound starch synthase in plants.Biotech Bull,2016,32(3):18-23.(in Chinese with English abstract)
[22]陈书强,杨丽敏,赵海新,杜晓东,周通,薛菁芳,单莉莉,王翠,张献国.穗粒肥用氮比例对寒地不同类型水稻产量和品质的影响.中国稻米.2017,23(4):151-156.Chen S Q,Yang L M,Zhao H X,Du X D,Zhou T,Xue JF,Shan L L,Wang C,Zhang X G.Effects of the spike-grain nitrogen ratio on yield and quality of different types of rice in cold region.China Rice,2017,23(4):151-156.(in Chinese)
[23]Kim W,Johnson J W,Graybosch R A,Gaines C S.Physicochemical properties and end-use quality of wheat starch as a function of waxy protein alleles.J Cereal Sci,2003,37(2):195-204.
[24]言普,沈文涛,黎小瑛,周鹏.植物hp RNA干扰载体构建的研究进展.生物技术通报,2013(9):7-12.Yan P,Shen W T,Li X Y,Zhou P.Progress in Construction of hp RNA vector for plant RNAi.Biotechnol Bull,2013(9):7-12.(in Chinese with English abstract)
[25]Sasaki T,Yasui T,Matsuki J.Effect of amylose content on gelatinization,retrogradation,and pasting properties of starches from waxy and nonwaxy wheat and their F1seeds.Cereal Chem,2000,77(1):58-63.
[26]Wang J C,Xu H,Zhu Y,Liu Q Q,Cai X L.Osb ZIP58,a basic leucine zipper transcription factor,regulates starch biosynthesis in rice endosperm//2012 National Conference on Plant Biology.2012:3453-3466.
[27]方先文,姜东,戴廷波,曹卫星.小麦淀粉组分的积累规律.江苏农业学报,2002,18(3):139-142.Fang X W,Jiang D,Dai T B,Cao W X.Accumulation regularity of starch components in wheat varieties.Jiangsu J Agric Sci,2002,18(3):139-142.(in Chinese with English abstract)
[28]Nishi A,Nakamura Y,Tanaka N,Satoh H.Organ Specificity of isoforms of starch branching enzyme(Q-enzyme)in rice.Plant Cell Physiol,1992,33(7):985-991.
[29]Satoh H,Nishi A,Yamashita K,Takemoto Y.Alteration of the structural properties of starch components by the lack of an isoforms of starch branching enzyme in rice seeds.J Biol Chem,1993,268(25):19084-19091.
[30]Kawasaki T,Mizuno K,Baba T,Shimada H.Molecular analysis of the gene encoding a rice starch branching enzyme.Mol Gen Genet,1993,237(1):10-16.
[31]Guan H P,Preiss J.Differentiation of the properties of the branching isozymes from maize(Zea mays).Plant Physiol,1993,102(4):1269-1273.
[32]Morell M K,Blennow A,Kosar-Hasherni B,Samuel MS.Differential expression and Properties of starch branching enzyme isoforms in developing wheat endosperm.Plant Physiol,1997,113(1):201-208.
[33]Satoh H,Nishi A,Yamashita K,Takemoto Y,Tanaka Y,Hosaka Y,Sakurai A,Fujita N,Nakamura Y.Starch-branching enzyme I deficient mutation specifically affects the structure and properties of starch in rice endosperm.Plant Physi,2003,133:1111-1121.
[34]Utsumi Y,Utsumi C,Sawada T,Fujita N,Nakamura Y.Functional diversity of isoamylase oligomers:The ISA1homo-oligomer is essential for amylopectin biosynthesis in rice endosperm.Plant Physiol,2011,156(1):61-77.
[35]Kubo A,Rahman S,Utsumi Y.Complementation of sugary-1phenotype in rice endosperm with the wheat isoamy-lase1gene supports a direct role forisoamylase1 in amylopectin biosynthesis.Plant Physiol,2005,137(1):43-56.
[36]钟连进,董虎,蔡小波,封言柠,任凭,程方民.控制水稻胚乳淀粉合成代谢若干关键酶基因对花后高温的响应表达.应用生态学报,2012,23(3):745-750.Zhong L J,Dong H,Cai X B,Feng Y N,Ren P,Cheng FM.Gene expression of the key enzymes controlling starch synthesis and metabolism in rice grain endosperm under effects of high temperature after anthesis.Chin JAppl Ecol,2012,23(3):745-750.(in Chinese with English abstract)
[37]钟连进,程方民,张国平,孙宗修.灌浆结实期不同温度下早籼稻米淀粉链长分布与结构特征差异分析.中国农业科学,2005,38(2):272-276.Zhong L J,Cheng F M,Zhang G P,Sun Z X.Differences in starch chain length distribution and structure characteristics of early-indica rice under different temperature treatments during grain-filling.Sci Agric Sin,2005,38(2):272-276.(in Chinese with English abstract)
[38]Delatte T,Trevisan M,Parker M L,Zeeman S C.Arabidopsis mutants Atisa1 and Atisa2 have identical phenotypes and lack the same multimeric isoamylase,which influences the branch point distribution of amylopectin during starch synthesis.Plant J,2005,41(6):815-830.