小麦强弱势粒胚乳淀粉体和蛋白体发育及物质积累研究
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摘要
为探明小麦强弱势粒发育的差异,以大穗型小麦品种宁麦13为材料,采用半薄切片、组织染色和生理生化测定等方法比较研究了小麦强弱势粒胚乳淀粉体和蛋白体的发育及物质积累过程。结果表明,在小麦颖果发育过程中,强势粒的形态大小始终大于弱势粒,在花后3～11d表现尤为明显;与弱势粒相比,强势粒的胚乳尤其是背部胚乳中淀粉体和蛋白体的相对面积较大,充实度较高,胚乳质地更致密;强势粒的B型淀粉粒多于弱势粒,强、弱势粒的淀粉粒径分布均呈单峰型,强势粒中0～2μm的淀粉粒数显著高于弱势粒,在其他粒径范围两者无显著差异;与弱势粒相比,强势粒中总淀粉、蛋白质、醇溶蛋白、谷蛋白和支链淀粉含量均较高,而直链淀粉含量较低。由此可见,小麦弱势粒发育不良与胚乳淀粉体和蛋白体发育及物质积累密切相关。
In order to clarify the developmental differences of endosperm between superior and inferior grains,Ningmai 13,a big-spike wheat variety,was used as the material to investigate amyloid and protein development and their accumulation in endosperm between superior and inferior wheat grains by the methods of semi-thin sections,tissue staining and physiological and biochemical determination.The main results were as follows,during development of the wheat grains,the size of superior grains were always bigger than that of inferior grains,especially in the period of 3to 11 days after flowering.Compared with inferior grains,superior grains,especially in the back endosperm,have larger relative areas of the amyloid and the protein,higher grain filling degree,and denser endosperm texture.The number of B-starch granules in superior grains is greater than those in inferior grains,and the starch size of superior and inferior grains showed unimodal distribution.Within the particle size range of 0-2μm,the proportion of starch granules of superior grains were significantly higher than that of inferior grains,but within other ranges,there was no significant differences.Compared with inferior grains,su-perior grains have higher content of the total starch,protein,alcohol soluble protein,gluten and amylopectin,but lower amylose content.The results above showed that poor development of inferior grains was relative to amyloid and protein development and their accumulation.
In order to clarify the developmental differences of endosperm between superior and inferior grains,Ningmai 13,a big-spike wheat variety,was used as the material to investigate amyloid and protein development and their accumulation in endosperm between superior and inferior wheat grains by the methods of semi-thin sections,tissue staining and physiological and biochemical determination.The main results were as follows,during development of the wheat grains,the size of superior grains were always bigger than that of inferior grains,especially in the period of 3to 11 days after flowering.Compared with inferior grains,superior grains,especially in the back endosperm,have larger relative areas of the amyloid and the protein,higher grain filling degree,and denser endosperm texture.The number of B-starch granules in superior grains is greater than those in inferior grains,and the starch size of superior and inferior grains showed unimodal distribution.Within the particle size range of 0-2μm,the proportion of starch granules of superior grains were significantly higher than that of inferior grains,but within other ranges,there was no significant differences.Compared with inferior grains,su-perior grains have higher content of the total starch,protein,alcohol soluble protein,gluten and amylopectin,but lower amylose content.The results above showed that poor development of inferior grains was relative to amyloid and protein development and their accumulation.
引文
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[9]徐云姬,李银银,钱希旸,等.三种禾谷类作物强、弱势粒淀粉粒形态与粒度分布的比较[J].作物学报,2016,42(1):71.XU Y J,LI Y Y,QIAN X Y,et al.Comparison of starch granule morphology and size distribution in superior and inferior grains of three cereal crops[J].Acta Agronomica Sinica,2016,42(1):71.
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[13]梁太波,尹燕枰,蔡瑞国,等.大穗型小麦品种强、弱势籽粒淀粉积累和相关酶活性的比较[J].作物学报,2008,34(1):155.LIANG T B,YIN Y P,CAI R G,et al.Starch accumulation and related enzyme activities in superior and inferior grains of large spike wheat[J].Acta Agronomica Sinica,2008,34(1):155.
[14]WANG LLYU X R,YANG Y,et al.Morphology and physicochemical properties of starch inwheatsuperior and inferior grains[J].Starch-St9rke,2017,70(30):1700135.
[15]ZHU T,JACKSON DS,WEHLINGRL,et al.Comparison of amylosedetermination methods and the development of a dualwavelengthiodine binding technique[J].Cereal Chemistry,2008,85(1):51.
[16]郑先哲.稻米食味值测定及干燥品质的研究[J].农业机械学报,2000,31(4):54.ZHENG X Z.Calculation of taste value and study on drying quality of rice[J].Transactions of the Chinese Society for Agricultural Machinery,2000,31(4):54.
[17]XIONGF,YU X R,ZHOU L,et al.Effect of nitrogen application at the booting stage on wheat progenyseed germination and seedling growth[J].Journal of Plant Studies,2013,2(2):159.
[18]YU X R,YU H,ZHANG J,et al.Comparison of endosperm starch granule development and physicochemical properties of starches from waxy and non-waxy wheat[J].International Journal of Food Properties,2015,18(11):2414.
[19]XIONGF,YU X R,ZHOUL,et al.The influence of nitrogen on the development and accumulation of protein bodies in the developing endosperm of wheat caryopses[J].Molecular Biology Reports,2014,41(2):690.
[20]郭艳艳.不同冬小麦品种籽粒胚乳增殖和灌浆对粒重的影响[J].麦类作物学报,2018,38(1):88.GUO Y Y.The effect of different winter wheat cultivars on the grain weight of seed endosperm proliferation and grouting[J].Journal of Triticeae Crops,2018,38(1):88.
[21]顾蕴洁,王忠,吴月萍.小麦胚乳细胞的分离及其淀粉体的计数[J].植物生理学通讯,1994(3):211.GU Y J,WANG Z,WU Y P.Isolation of endosperm cells and calculation of their starch grains in wheat plants[J].Plant Physiology Communications,1994(3):211.
[22]刘智.关于稻麦胚乳细胞发育的研究[D].扬州:扬州大学,2012:167.LIU Z.Study on the development of rice endosperm cells[D].Yangzhou:Yangzhou University,2012:167.
[23]DOUGLAS C D,KOU T M,FELKER F C.Enzymes of sucrose and hexose metabolism in developing kernels of two inbreds of maize[J].Plant Physiology,1988,86(4):1015.
[24]谭秀山,毕建杰,王金花,等.冬小麦不同穗位籽粒淀粉粒差异及其与粒重的相关性[J].作物学报,2012,38(10):1923.TAN X S,BI J J,WANG J H,et al.Differences of starch granules in grans from different spikelet positions and their correlation with grain weight in winter wheat[J].Acta Agronomica Sinica,2012,38(10):1923.
[25]RAMESH M,MITCHELL J R,JUMEL K,et al.What is the true amylose content of rice starch[C]//Gums and Stabilisers for the Food Industry,Woodhead Publishing,2000:76.
[26]CHOPE GA,WAN Y,PENSON SP,et al.Effects of genotype,season,and nitrogennutrition on gene expression and protein accumulation in wheat grain[J].Journal of Agricultural and Food Chemistry,2014,62(19):4405.
[27]TOSI P.Traffcking and deposition of prolamins in wheat[J].Journal of Cereal Science,2012,56(1):85.
[28]WAN Y,SHEWRY P R,HAWKESFORD M J.A novel family ofγ-gliadingenesis highly regulated by nitrogen supply in developing wheat grain[J].Journal of Experimental Botany,2013,64(1):166.
[29]SHEWRY P R,HALFORD N G,TATHAM A S,et al.The high molecular weight subunits of wheat gluteninand their role in determining wheatprocessingproperties[J].Advances in Food and Nutrition Research,2003,45(3):223.
[2]SABELLI P A,LARKINS B A.The development of endosperm in grasses[J].Plant Physiology,2009,149(1):14.
[3]王蔚华,郭文善,方明荃,等.小麦籽粒胚乳细胞增殖及物质充实动态[J].作物学报,2003,29(5):780.WANG W H,GUO W S,FANG M Q,et al.Endosperm cell proliferating and grain filling dynamics in wheat[J].Acta Agronomica Sinica,2003,29(5):780.
[4]封超年,郭文善,施劲松,等.小麦花后高温对籽粒胚乳细胞发育及粒重的影响[J].作物学报,2000,26(4):400.FENG C N,GUO W S,SHI J S,et al.Effect of high temperature after anthesis on endosperm cell development and grain weight in wheat[J].Acta Agronomica Sinica,2000,26(4):400.
[5]王忠,顾蕴洁,李卫芳,等.小麦胚乳发育及其养分输入的途径[J].作物学报,1998,24(5):536.WANG Z,GU Y J,LI W F,et al.Development of wheat wndosperm and the pathway of nutrient entering the endosperm[J].Acta Agronomica Sinica,1998,24(5):536.
[6]荆彦平,刘大同,李栋梁,等.小麦内胚乳细胞及其淀粉体的发育[J].麦类作物学报,2013,33(4):820.JING Y P,LIU D T,LI D L,et al.Development of starch endosperm cells and amyloplasts in wheat[J].Journal of Triticeae Crops,2013,33(4):820.
[7]CHEN X,LI B,SHAO S,et al.Accumulation characteristic of protein bodies in different regions of wheat endosperm under drought stress[J].Journal of Integrative Agriculture,2016,15(12):2927.
[8]YU X R,LI B,WANG L L,et al.Effect of drought stress on the development of endosperm starch granules and the composition and physicochemical properties of starches from soft and hard wheat[J].Journal of the Science of Food and Agriculture,2016,96(8):2746.
[9]徐云姬,李银银,钱希旸,等.三种禾谷类作物强、弱势粒淀粉粒形态与粒度分布的比较[J].作物学报,2016,42(1):71.XU Y J,LI Y Y,QIAN X Y,et al.Comparison of starch granule morphology and size distribution in superior and inferior grains of three cereal crops[J].Acta Agronomica Sinica,2016,42(1):71.
[10]靳德明.杂交水稻强弱势颖花异步灌浆结实特性及其生理机制研究[D].武汉:华中农业大学,2002:43.JIN D M.Study on the strong characteristics and physiological mechanism of the hybrid rice with the strong and weak and the asynchronous grout[D].Wuhan:Huazhong Agricultural University,2002:43.
[11]刘智,王玲玲,张二金,等.不同类型专用小麦品种胚乳发育的比较研究[J].麦类作物学报,2011,31(1):70.LIU Z,WANG L L,ZHANG E J,et al.Study on endosperm development between wheat cultivars for different users[J].Journal of Triticeae Crops,2011,31(1):70.
[12]孟秀蓉,熊飞,孔妤,等.强、中、弱筋小麦籽粒中淀粉、蛋白质积累和淀粉体发育的比较[J].作物学报,2009,35(5):962.MENG X R,XIONG F,KONG Y,et al.Comparison of starch,protein accumulation and amyloplastdevelopment inwheatcultivars with strong,medium,and weak gluten[J].Acta Agronomica Sinica,2009,35(5):962.
[13]梁太波,尹燕枰,蔡瑞国,等.大穗型小麦品种强、弱势籽粒淀粉积累和相关酶活性的比较[J].作物学报,2008,34(1):155.LIANG T B,YIN Y P,CAI R G,et al.Starch accumulation and related enzyme activities in superior and inferior grains of large spike wheat[J].Acta Agronomica Sinica,2008,34(1):155.
[14]WANG LLYU X R,YANG Y,et al.Morphology and physicochemical properties of starch inwheatsuperior and inferior grains[J].Starch-St9rke,2017,70(30):1700135.
[15]ZHU T,JACKSON DS,WEHLINGRL,et al.Comparison of amylosedetermination methods and the development of a dualwavelengthiodine binding technique[J].Cereal Chemistry,2008,85(1):51.
[16]郑先哲.稻米食味值测定及干燥品质的研究[J].农业机械学报,2000,31(4):54.ZHENG X Z.Calculation of taste value and study on drying quality of rice[J].Transactions of the Chinese Society for Agricultural Machinery,2000,31(4):54.
[17]XIONGF,YU X R,ZHOU L,et al.Effect of nitrogen application at the booting stage on wheat progenyseed germination and seedling growth[J].Journal of Plant Studies,2013,2(2):159.
[18]YU X R,YU H,ZHANG J,et al.Comparison of endosperm starch granule development and physicochemical properties of starches from waxy and non-waxy wheat[J].International Journal of Food Properties,2015,18(11):2414.
[19]XIONGF,YU X R,ZHOUL,et al.The influence of nitrogen on the development and accumulation of protein bodies in the developing endosperm of wheat caryopses[J].Molecular Biology Reports,2014,41(2):690.
[20]郭艳艳.不同冬小麦品种籽粒胚乳增殖和灌浆对粒重的影响[J].麦类作物学报,2018,38(1):88.GUO Y Y.The effect of different winter wheat cultivars on the grain weight of seed endosperm proliferation and grouting[J].Journal of Triticeae Crops,2018,38(1):88.
[21]顾蕴洁,王忠,吴月萍.小麦胚乳细胞的分离及其淀粉体的计数[J].植物生理学通讯,1994(3):211.GU Y J,WANG Z,WU Y P.Isolation of endosperm cells and calculation of their starch grains in wheat plants[J].Plant Physiology Communications,1994(3):211.
[22]刘智.关于稻麦胚乳细胞发育的研究[D].扬州:扬州大学,2012:167.LIU Z.Study on the development of rice endosperm cells[D].Yangzhou:Yangzhou University,2012:167.
[23]DOUGLAS C D,KOU T M,FELKER F C.Enzymes of sucrose and hexose metabolism in developing kernels of two inbreds of maize[J].Plant Physiology,1988,86(4):1015.
[24]谭秀山,毕建杰,王金花,等.冬小麦不同穗位籽粒淀粉粒差异及其与粒重的相关性[J].作物学报,2012,38(10):1923.TAN X S,BI J J,WANG J H,et al.Differences of starch granules in grans from different spikelet positions and their correlation with grain weight in winter wheat[J].Acta Agronomica Sinica,2012,38(10):1923.
[25]RAMESH M,MITCHELL J R,JUMEL K,et al.What is the true amylose content of rice starch[C]//Gums and Stabilisers for the Food Industry,Woodhead Publishing,2000:76.
[26]CHOPE GA,WAN Y,PENSON SP,et al.Effects of genotype,season,and nitrogennutrition on gene expression and protein accumulation in wheat grain[J].Journal of Agricultural and Food Chemistry,2014,62(19):4405.
[27]TOSI P.Traffcking and deposition of prolamins in wheat[J].Journal of Cereal Science,2012,56(1):85.
[28]WAN Y,SHEWRY P R,HAWKESFORD M J.A novel family ofγ-gliadingenesis highly regulated by nitrogen supply in developing wheat grain[J].Journal of Experimental Botany,2013,64(1):166.
[29]SHEWRY P R,HALFORD N G,TATHAM A S,et al.The high molecular weight subunits of wheat gluteninand their role in determining wheatprocessingproperties[J].Advances in Food and Nutrition Research,2003,45(3):223.