糜子籽粒形成过程中蛋白质、淀粉积累与相关合成酶特性
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摘要
为探究糜子灌浆过程中籽粒蛋白质及淀粉积累规律,选用糯性和粳性糜子品种各2个,分别为‘龙黍21号’、‘晋黍5号’和‘榆糜2号’、‘陇糜8号’,分析淀粉合成相关酶活性的动态特性。结果表明:糜子开花后,粳糯品种籽粒脂肪含量先增加后减少,而粗蛋白含量不断减少,其中谷蛋白含量最高。各氨基酸中,谷氨酸、脯氨酸和亮氨酸含量最高。淀粉含量变化呈S型增加,积累速率先增加后减小。4种淀粉合成相关酶(ADP-葡萄糖焦磷酸化酶(AGPP)、可溶性淀粉合成酶(SSS)、颗粒结合性淀粉合成酶(GBSS)以及淀粉分支酶(Q酶))活性显示出单峰曲线变化。粳性品种中SSS酶主要调控直链、支链及总淀粉积累,Q酶主要调控总淀粉和直链淀粉,AGPP酶参与直链淀粉的积累;而糯性品种直链、支链及总淀粉积累主要受到SSS和Q酶的调控。
The dynamics of protein and starch accumulation and the activities of starch synthesis related enzymes during grain filling of proso millet were investigated in this study.Two waxy varieties(‘Longshu 21'and ‘Jinshu 5') and two non-waxy varieties(‘Yumi 2'and ‘Longmi 8') were taken as study materials.The results showed that the content of fat increased firstly and then decreased after anthesis.However,the content of crude protein continuously decreased,among which gluten content was the highest.Proso millet grain was found rich in glutamic acid,proline and leucine.And the accumulation of total starch,amylose and amylopectin showed a trend of S-type growth and their accumulation rates increased firstly and decreased later.The activity of AGPP,SSS,GBSS and Q enzyme showed a single peak curve during the process of grain filling.For non-waxy varieties,the accumulation rate of total starch was correlated with the activity of SSS and Q enzyme,its amylose was significantly affected by the AGPP,SSS and Q enzyme.SSS was pivotal in amylopectin accumulation of non-waxy varieties.While the activity of SSS and Q enzyme played key roles in the process of all accumulation of waxy varieties.
The dynamics of protein and starch accumulation and the activities of starch synthesis related enzymes during grain filling of proso millet were investigated in this study.Two waxy varieties(‘Longshu 21'and ‘Jinshu 5') and two non-waxy varieties(‘Yumi 2'and ‘Longmi 8') were taken as study materials.The results showed that the content of fat increased firstly and then decreased after anthesis.However,the content of crude protein continuously decreased,among which gluten content was the highest.Proso millet grain was found rich in glutamic acid,proline and leucine.And the accumulation of total starch,amylose and amylopectin showed a trend of S-type growth and their accumulation rates increased firstly and decreased later.The activity of AGPP,SSS,GBSS and Q enzyme showed a single peak curve during the process of grain filling.For non-waxy varieties,the accumulation rate of total starch was correlated with the activity of SSS and Q enzyme,its amylose was significantly affected by the AGPP,SSS and Q enzyme.SSS was pivotal in amylopectin accumulation of non-waxy varieties.While the activity of SSS and Q enzyme played key roles in the process of all accumulation of waxy varieties.
引文
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[13] 杨清华,邱军,李海,杨天育,程炳文,赵敏,刘国庆,高小丽,冯佰利.糜子育成品种农艺、产量及品质性状综合鉴定与评价[J].中国农业科学,2017,50(23):4530-4544Yang Q H,Qiu J,Li H,Yang T Y,Cheng B W,Zhao M,Liu G Q,Gao X L,Feng B L.Comprehensive evaluation of agronomic,yield and quality traits of broomcorn millet (Panicum miliaceum L) cultivars[J].Scientia Agricultura Sinica,2017,50(23):4530-4544 (in Chinese)
[14] 柴岩.糜子(黄米)的营养和生产概况[J].粮食加工,2009,34(4):90-91Chai Y.Nutrition and production overview of proso millet (yellow rice)[J].Grain Processing,2009,34(4):90-91 (in Chinese)
[15] Zarnowski R,Suzuki Y.Expedient Soxhlet extraction of resorcinolic lipids from wheat grains[J].Journal of Food Composition & Analysis,2004,17(5):649-663
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[18] Yamashita S,Baba N,Moriyama H.Application of dual-wavelength spectrophotometry to determination of amylose and amylopectin contents[J].Nippon Shokuhin Kagaku Kogaku Kaishi,2011,40(5):365-369
[19] Nakamura Y,Yuki K,Park S Y,Ohya T.Carbohydrate metabolism in the developing endosperm of rice grains[J].Plant & Cell Physiology,1989,30(6):833-839
[20] 赵法茂,齐霞,肖军,王宪泽.测定淀粉分支酶活性方法的改进[J].植物生理学报,2007,43(6):1167-1169Zhao F M,Qi X,Xiao J,Wang X Z.Improvemed method for determining starch branching enzyme activity[J].Plant Physiology Journal,2007,43(6):1167-1169 (in Chinese)
[21] 田翔,乔治军,田琴,曹晓宁,王君杰,刘思辰.气相色谱-质谱法定性定量测定糜子中的脂肪酸[J].食品科学,2016,37(14):128-131Tian X,Qiao Z J,Tian Q,Cao X N,Wang J J,Liu S C.Identification and quantitation of fatty acids in Panicum miliaceum L by gas chromatography-mass spectrometry[J].Food Science,2016,37(14):128-131 (in Chinese)
[22] 许振柱,周广胜,植物氮代谢及其环境调节研究进展[J],应用生态学报,2004,15(3):511-516Xu Z Z,Zhou G S.Research advance in nitrogen metabolism of plant and its environmental regulation[J].The Journal of Applied Ecology,2004,15(3):511-516 (in Chinese)
[23] Desai R M,Bhatia C R.Nitrogen uptake and nitrogen harvest index in durum wheat cultivars varying in their grain protein concentration[J].Euphytica,1978,27(2):561-566
[24] 张智猛,戴良香,胡昌浩,董树亭,王空军.氮素对玉米淀粉累积及相关酶活性的影响[J].作物学报,2005,31(7):956-963Zhang Z M,Dai L X,Hu C H,Dong S T,Wang K J.Effect of nitrogen on starch accumulation and related enzyme activities maize (Zea mays) [J].Acta Agronomica Sinica,2005,31(7):956-963 (in Chinese)
[25] 林红.甜玉米籽粒营养成分动态变化研究[J].黑龙江农业科学,2011(2):5-7Lin H.Study on dynamic changes of the nutritional component of sweet corn grain[J].Heilongjiang Agricultural Sciences,2011(2):5-7 (in Chinese)
[26] Egli D B,Bruening W P.Accumulation of nitrogen and dry matter by soybean seeds with genetic differences in protein concentration[J].Crop Science,2007,47(1):359-366
[27] 陈洁.水稻籽粒淀粉和蛋白质积累模拟模型研究[D].南京:南京农业大学,2011Chen J.Study on rice grain starch and protein accumulation simulationmodel[D].Nanjing:Nanjing Agricultural University,2011 (in Chinese)
[28] 揭厚胜.不同直链淀粉含量的水稻品种籽粒淀粉和蛋白质积累特性的研究[D].雅安:四川农业大学,2004Jie H S.Study on characteristics of starch and protein accumulation in different amylose content rice varieties[D].Yaan:Sichuan Agricultural University,2004 (in Chinese)
[29] 谭彩霞,封超年,郭文善,朱新开,李春燕,彭永欣.不同小麦品种籽粒淀粉合成酶基因的表达及其与淀粉积累的关系[J].麦类作物学报,2011,31(6):1063-1070Tan C X,Feng C N,Guo W S,Zhu X K,Li C Y,Peng Y X.Difference in expression of starch synthase gene and starch synthesis in the grains of different wheat cultivars[J].Journal of Triticeae Crops,2011,31(6):1063-1070 (in Chinese)
[30] 张艳平,苏旺,高小丽,雷锦银,高金锋,王鹏科,冯佰利.不同覆盖栽培糜子籽粒灌浆期淀粉酶活性变化[J].中国农业大学学报,2014,19(1):51-58Zhang Y P,Su W,Gao X L,Lei J Y,Gao J F,Wang P K,Feng B L.Dynamic changes of starch synthesis enzymes activities of Panicum miliaceum L grain during the filling stage under different mulching cultivation[J].Journal of China Agricultural University,2014,19(1):51-58 (in Chinese)
[31] 韩浩坤.糜子籽粒形成过程中淀粉及蛋白质积累特性研究[D].杨凌:西北农林科技大学,2017Han H K.The starch and protein accumulation characteristics during the development of proso millet[D].Yangling:Northwest A & F University,2017 (in Chinese)
[2] Wang E T,Wang J J,Zhu X D,Hao W,Wang L Y,Li Q,Zhang L X,He W,Lu B R,Lin H X,Ma H,Zhang G Q,He Z H.Control of rice grain-filling and yield by a gene with a potential signature of domestication[J].Nature Genetics,2008,40(11),1370-1374
[3] Takai T,Fukuta Y,Shiraiwa T,Horie T.Time-related mapping of quantitative trait loci controlling grain-filling in rice (Oryza sativa L)[J].Journal of Experimental Botany,2005,56(418):2107-2118
[4] 于凤瑶,辛秀君,张代军,周顺启,邱红梅.大豆籽粒干物质、脂肪和蛋白质的动态积累规律研究[J].农业现代化研究,2009,30(5):637-640Yu F Y,Xin X J,Zhang D J,Zhou S Q,Qiu H M.Dynamic accumulation of dry matter,oil and protein in soybean seed[J].Research of Agricultural Modernization,2009,30(5):637-640 (in Chinese)
[5] 徐一兰,官春云,谭太龙,余龙喜.油菜种子形成中含油量与其合成相关酶活性的变化及其相关性[J].作物学报,2008,34(10):1854-1857Xu Y L,Guan C Y,Tan T L,Yu L X.Changes of oil content and oil biosynthesis-related enzymes activities and their correlation during seed formation in Brassica napus[J].Acta Agronomica Sinica,2008,34(10):1854-1857 (in Chinese)
[6] 赵勇,张树华,王杰,杨学芳,李晓云,鞠晓影,杨学举.小麦籽粒灌浆期蛋白质组分含量变化规律的研究[J].河北农业大学学报,2016,39(5):8-11Zhao Y,Zhang S H,Wang J,Yang X F,Li X Y,Ju X Y,Yang X J.Investigation to the pattern for dynamic diversification of protein-fraction contents during grain development in wheat[J].Journal of Agricultural University of Hebei,2016,39(5):8-11 (in Chinese)
[7] 周国勤,张应香,李宇峰,李明慧,李刚.信阳234的籽粒灌浆特性及蛋白质、淀粉积累动态研究[J].山东农业科学,2008,1(1):51-54Zhou G Q,Zhang Y X,Li Y F,Li M H,Li G.Study of grain filling characteristics and accumulation dynamics of protein and starch in grains of wheat cultivar Xinyang 234[J].Shandong Agricultural Sciences,2008,1(1):51-54 (in Chinese)
[8] 方先文,姜东,戴廷波,曹卫星.小麦淀粉组分的积累规律[J].江苏农业学报,2002,18(3):139-142Fang X W,Jiang D,Dai Y B,Cao W X.Accumulation regularity of starch components in wheat varieties[J].Jiangsu Journal of Agricultural Sciences,2002,18(3):139-142 (in Chinese)
[9] Ballicora M A,Iglesias A A,Preiss J.ADP-Glucose pyrophosphorylase:A regulatory enzyme for plant starch synthesis[J].Photosynthesis Research,2004,79(1):1
[10] 王自布,黄燕芬,吴坤,任翠娟,姜金仲.籽粒淀粉合成酶与淀粉合成关系的研究进展[J].生物技术进展,2013,1(5):336-341Wang Z B,Huang Y F,Wu K,Ren C J,Jiang J Z.Progress on correlation between starch synthase and starch synthesis in the grain[J].Current Biotechnology,2013(5):336-341 (in Chinese)
[11] Qu Y,Su W,Zhang P P,Li C.Effects of different water harvesting on soil water,growth and yield of the proso millet (Panicum miliaceum L.) in a semiarid region of Northwest China[J].Journal of Agricultural Science,2012,4(9):106-113
[12] Zhang Y Y,Han H K,Zhang D Z,Li J,Gong X W,Feng B L,Xue Z H,Yang P.Effects of ridging and mulching combined practices on proso millet growth and yield in semi-arid regions of China[J].Field Crops Research,2017,213(1):65-74
[13] 杨清华,邱军,李海,杨天育,程炳文,赵敏,刘国庆,高小丽,冯佰利.糜子育成品种农艺、产量及品质性状综合鉴定与评价[J].中国农业科学,2017,50(23):4530-4544Yang Q H,Qiu J,Li H,Yang T Y,Cheng B W,Zhao M,Liu G Q,Gao X L,Feng B L.Comprehensive evaluation of agronomic,yield and quality traits of broomcorn millet (Panicum miliaceum L) cultivars[J].Scientia Agricultura Sinica,2017,50(23):4530-4544 (in Chinese)
[14] 柴岩.糜子(黄米)的营养和生产概况[J].粮食加工,2009,34(4):90-91Chai Y.Nutrition and production overview of proso millet (yellow rice)[J].Grain Processing,2009,34(4):90-91 (in Chinese)
[15] Zarnowski R,Suzuki Y.Expedient Soxhlet extraction of resorcinolic lipids from wheat grains[J].Journal of Food Composition & Analysis,2004,17(5):649-663
[16] 邓妙,魏益民.蛋白质组分的连续累进提取分析法[J].西北农林科技大学学报:自然科学版,1989,1(1):110-113Deng M,Wei Y M.Successive extraction and analysis of protein fractions[J].Journal of Northwest A&F University:Natural Science Edition,1989,1(1):110-113 (in Chinese)
[17] GB/T5009.124—2003.食品中氨基酸的测定[S].北京:中国标准出版社,2003GB/T5009.124—2003.Determination of amino acids in food[S].Beijing:Standards Press of China,2003
[18] Yamashita S,Baba N,Moriyama H.Application of dual-wavelength spectrophotometry to determination of amylose and amylopectin contents[J].Nippon Shokuhin Kagaku Kogaku Kaishi,2011,40(5):365-369
[19] Nakamura Y,Yuki K,Park S Y,Ohya T.Carbohydrate metabolism in the developing endosperm of rice grains[J].Plant & Cell Physiology,1989,30(6):833-839
[20] 赵法茂,齐霞,肖军,王宪泽.测定淀粉分支酶活性方法的改进[J].植物生理学报,2007,43(6):1167-1169Zhao F M,Qi X,Xiao J,Wang X Z.Improvemed method for determining starch branching enzyme activity[J].Plant Physiology Journal,2007,43(6):1167-1169 (in Chinese)
[21] 田翔,乔治军,田琴,曹晓宁,王君杰,刘思辰.气相色谱-质谱法定性定量测定糜子中的脂肪酸[J].食品科学,2016,37(14):128-131Tian X,Qiao Z J,Tian Q,Cao X N,Wang J J,Liu S C.Identification and quantitation of fatty acids in Panicum miliaceum L by gas chromatography-mass spectrometry[J].Food Science,2016,37(14):128-131 (in Chinese)
[22] 许振柱,周广胜,植物氮代谢及其环境调节研究进展[J],应用生态学报,2004,15(3):511-516Xu Z Z,Zhou G S.Research advance in nitrogen metabolism of plant and its environmental regulation[J].The Journal of Applied Ecology,2004,15(3):511-516 (in Chinese)
[23] Desai R M,Bhatia C R.Nitrogen uptake and nitrogen harvest index in durum wheat cultivars varying in their grain protein concentration[J].Euphytica,1978,27(2):561-566
[24] 张智猛,戴良香,胡昌浩,董树亭,王空军.氮素对玉米淀粉累积及相关酶活性的影响[J].作物学报,2005,31(7):956-963Zhang Z M,Dai L X,Hu C H,Dong S T,Wang K J.Effect of nitrogen on starch accumulation and related enzyme activities maize (Zea mays) [J].Acta Agronomica Sinica,2005,31(7):956-963 (in Chinese)
[25] 林红.甜玉米籽粒营养成分动态变化研究[J].黑龙江农业科学,2011(2):5-7Lin H.Study on dynamic changes of the nutritional component of sweet corn grain[J].Heilongjiang Agricultural Sciences,2011(2):5-7 (in Chinese)
[26] Egli D B,Bruening W P.Accumulation of nitrogen and dry matter by soybean seeds with genetic differences in protein concentration[J].Crop Science,2007,47(1):359-366
[27] 陈洁.水稻籽粒淀粉和蛋白质积累模拟模型研究[D].南京:南京农业大学,2011Chen J.Study on rice grain starch and protein accumulation simulationmodel[D].Nanjing:Nanjing Agricultural University,2011 (in Chinese)
[28] 揭厚胜.不同直链淀粉含量的水稻品种籽粒淀粉和蛋白质积累特性的研究[D].雅安:四川农业大学,2004Jie H S.Study on characteristics of starch and protein accumulation in different amylose content rice varieties[D].Yaan:Sichuan Agricultural University,2004 (in Chinese)
[29] 谭彩霞,封超年,郭文善,朱新开,李春燕,彭永欣.不同小麦品种籽粒淀粉合成酶基因的表达及其与淀粉积累的关系[J].麦类作物学报,2011,31(6):1063-1070Tan C X,Feng C N,Guo W S,Zhu X K,Li C Y,Peng Y X.Difference in expression of starch synthase gene and starch synthesis in the grains of different wheat cultivars[J].Journal of Triticeae Crops,2011,31(6):1063-1070 (in Chinese)
[30] 张艳平,苏旺,高小丽,雷锦银,高金锋,王鹏科,冯佰利.不同覆盖栽培糜子籽粒灌浆期淀粉酶活性变化[J].中国农业大学学报,2014,19(1):51-58Zhang Y P,Su W,Gao X L,Lei J Y,Gao J F,Wang P K,Feng B L.Dynamic changes of starch synthesis enzymes activities of Panicum miliaceum L grain during the filling stage under different mulching cultivation[J].Journal of China Agricultural University,2014,19(1):51-58 (in Chinese)
[31] 韩浩坤.糜子籽粒形成过程中淀粉及蛋白质积累特性研究[D].杨凌:西北农林科技大学,2017Han H K.The starch and protein accumulation characteristics during the development of proso millet[D].Yangling:Northwest A & F University,2017 (in Chinese)