糯大麦与非糯大麦颖果发育过程的解剖学与组织化学研究
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摘要
为了明确糯大麦与非糯大麦颖果发育的差异,该研究以糯大麦代表性品种(‘白青稞’、‘甘垦5号’)和非糯大麦代表性品种(‘扬农啤10号’和‘苏裸麦1号’)为材料,采用体视显微镜观察、组织化学染色、树脂半薄切片和光学显微镜观察等方法,比较研究了糯大麦与非糯大麦颖果及其胚乳淀粉体发育的过程。结果显示:(1)糯大麦和非糯大麦颖果的形态变化,以及鲜、干重和含水率等的变化规律基本一致,生长曲线均呈"S"型。(2)两类大麦颖果胚乳和果皮的I2/KI染色结果不同,糯大麦胚乳的I2/KI染色结果为红褐色,其果皮被染为蓝黑色,而非糯大麦胚乳和果皮均被I2/KI染成蓝黑色,表明糯大麦和非糯大麦的颖果果皮里直链淀粉含量都较高,且糯大麦胚乳内以支链淀粉为主,而非糯大麦的胚乳内以直链淀粉为主。(3)糯大麦胚乳淀粉体的出现时间早于非糯大麦,且其中小淀粉体的比例高于非糯大麦,淀粉体充实状况也好于非糯大麦。(4)与非糯大麦相比,糯大麦表观直链淀粉以及总淀粉含量较低,但可溶性糖含量较高。研究表明,糯大麦颖果生长规律与非糯大麦类似,但内含物淀粉的积累规律不同。
To explore the difference between waxy and non-waxy barley,we used two waxy barley and two non-waxy barley cultivars as materials in this paper.The development of caryopsis in waxy and non-waxy barley were investigated by stereomicroscopy observation,histochemical staining,resin semi-thin slicing and optical microscope observation.The results showed that:(1)the changes of fresh weight,dry weight and moisture content of waxy and non-waxy barley were basically consistent,and the growth curve was as‘S'.(2)The results of I2/KI staining for the endosperm and pericarp of two kinds of barley were different.The endosperm of waxy barley was stained red-brown by I2/KI and its pericarp was stained blue-black,but the endosperm and pericarp of waxy barley were both stained blue-black by I2/KI,and both waxy and nonwaxy barley have high amylose content in the pericarp.(3)The occurrence of waxy barley endosperm amyloplast was earlier than that of non-waxy barley,and the proportion of small amyloplast was higher than that of non-waxy barley.Also,the endosperm starch filling degree of waxy barley was better than that of non-waxy barley.(4)Compared with non-waxy barley,waxy barley showed lower apparent amylase and total starch content but higher soluble sugar content.The results show that the caryopsis growth lawof waxy barley is similar to that of non-waxy barley,but the accumulation law of the starch is different.
To explore the difference between waxy and non-waxy barley,we used two waxy barley and two non-waxy barley cultivars as materials in this paper.The development of caryopsis in waxy and non-waxy barley were investigated by stereomicroscopy observation,histochemical staining,resin semi-thin slicing and optical microscope observation.The results showed that:(1)the changes of fresh weight,dry weight and moisture content of waxy and non-waxy barley were basically consistent,and the growth curve was as‘S'.(2)The results of I2/KI staining for the endosperm and pericarp of two kinds of barley were different.The endosperm of waxy barley was stained red-brown by I2/KI and its pericarp was stained blue-black,but the endosperm and pericarp of waxy barley were both stained blue-black by I2/KI,and both waxy and nonwaxy barley have high amylose content in the pericarp.(3)The occurrence of waxy barley endosperm amyloplast was earlier than that of non-waxy barley,and the proportion of small amyloplast was higher than that of non-waxy barley.Also,the endosperm starch filling degree of waxy barley was better than that of non-waxy barley.(4)Compared with non-waxy barley,waxy barley showed lower apparent amylase and total starch content but higher soluble sugar content.The results show that the caryopsis growth lawof waxy barley is similar to that of non-waxy barley,but the accumulation law of the starch is different.
引文
[1]王忠,顾蕴洁,王敏,等.关于水稻胚乳淀粉体发育的研究[EB/OL].北京:中国科技论文在线[2012-04-09].http://www.paper.edu.cn/releasepaper/content/201204-121.WANG Z,GU Y J,WANG M,et al.Study on the development of rice endosperm starch[EB/OL].Pe King,Sciencepaper Online[2012-04-09].http://www.paper.edu.cn/releasepaper/content/201204-121.
[2]杨智敏,孔德媛,杨晓云,等.青稞籽粒淀粉含量的差异[J].麦类作物学报,2013,33(6):1 139-1 143.YANG Z M,KONG D Y,YANG X Y,et al.The difference of starch content in highland barley[J].Journal of Triticeae Crops,2013,33(6):1 139-1 143.
[3]LI W H,XIAO X L,ZHANG W H,et al.Compositional,morphological,structural and physicochemical properties of starches from seven naked barley cultivars grown in China[J].Food Research International,2014,58(4):7-14.
[4]郑彦坤,曾德二,魏和平,等.水稻胚乳组织的结构观察[J].中国水稻科学,2017,31(1):91-98.ZHENG Y K,ZENG D E,WEI H P,et al.Structure observation of rice endosperm tissue[J].Chin.J.Rice Sci.,2017,31(1):91-98.
[5]王忠.水稻的开花与结实—水稻生殖器官发育图谱[M].北京:科学出版社,2015.
[6]李祥栋,石明,魏心元.禾谷类作物胚乳淀粉合成及Waxy基因研究进展[J].中国农学通报,2015,31(12):181-186.LI X D,SHI M,WEI X Y.Research progress of starch synthesis and Waxy gene in cereal crops[J].Chinese Agriculture Bulletin,2015,31(12):181-186.
[7]BERGER F.Endosperm development[J].Current Opinion in Plant Biology,1999,2(1):28-32.
[8]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):818-824.JIN Y P,LIU D T,LI D L,et al.The development of endosperm cells and their amyloid in wheat[J].Journal of Triticeae Crops,2013,33(4):818-824.
[9]VAN DONGEN J T,ROEB G W,DAUTZENBERG M,et al.Phloem import and storage metabolism are highly coordinated by the low oxygen concentrations within developing wheat seeds[J].Plant Physiology,2004,135(3):1 809-1 821.
[10]刘美,张凤,杨翠翠,等.小麦种子萌发早期淀粉降解关键酶活性及基因表达量研究[J].山东农业科学,2014,46(9):39-45.LIU M,ZHANG F,YANG C C,et al.Study on key enzyme activity and gene expression in early starch degradation of wheat seed germination[J].Shandong Agricultural Sciences,2014,46(9):39-45.
[11]SUH D S,VERHOEVEN T,DENYER K,et al.Characterization of Nubet and Franubet barley starches[J].Carbohydrate Polymers,2004,56(1):85-93.
[12]TANG H,MITSUNAGA T,KAWAMURA Y.Relationship between functionality and structure in barley starches[J].Carbohydrate Polymers,2004,57(2):145-152.
[13]高俊凤.植物生理学实验指导[M].北京:高等教育出版社,2006.
[14]张艺祎,李云霞,骆亮,等.低温储藏期间3种独蒜兰碳水化合物含量变化研究[EB/OL].北京:中国科技论文在线[2016-12-29].http://www.paper.edu.cn/releasepaper/content/201701-56.ZHANG Y Y,LI Y X,LUO L,et al.Study on the change of carbohydrate content of 3kinds of allicin in low temperature storage[EB/OL].PeKing:Sciencepaper Online[2016-12-29].http://www.paper.edu.cn/releasepaper/content/201701-56.
[15]余徐润,周亮,荆彦平,等.Image-Pro Plus软件在小麦淀粉粒显微图像分析中的应用[J].电子显微学报,2013,32(4):344-351.YU X R,ZHOU L,JING Y P,et al.Image-Pro Plus software used in wheat starch granule micrograph analysis[J].Journal of Chinese Electron Microscopy Society,2013,32(4):344-351.
[16]朱帆,徐广文,丁文平.表观直链淀粉含量对小麦淀粉凝胶流变特性影响的研究[J].食品科学,2008,29(2):43-47.ZHU F,XU G W,DING W P.Study on the effect of amylose content on the rheological properties of wheat starch gel[J].Food Science,2008,29(2):43-47.
[17]李栋梁,荆彦平,李小刚,等.高粱胚乳细胞与母体组织发育关系的研究[J].中国农业科学,2014,47(17):3 336-3 347.LI D L,JING Y P,LI X G,et al.Study on the relationship between sorghum endosperm cells and maternal tissue development[J].Scientia Agricultura Sinica,2014,47(17):3 336-3 347.
[18]王丹.夏橙果实成熟过程中果皮色泽变化的规律研究[D].武汉:华中农业大学,2016:1-2.
[19]荆彦平,李栋梁,刘大同,等.两个玉米品种胚乳发育的比较[J].玉米科学,2014,22(1):79-85.JING Y P,LI D L,LIU D T,et al.Comparison of endosperm development of two maize varieties[J].Journal of Maize Sciences,2014,22(1):79-85.
[20]苗红霞,孙佩光,张凯星,等.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[J].生物技术通报,2016,32(3):18-23.MIAO H X,SUN P G,ZHANG K X,et al.Research progress of the expression regulation mechanism of plant granule combined with starch synthase(GBSS)gene[J].Biotechnology Bulletin,2016,32(3):18-23.
[21]KRISHNAN H.B.,CHEN M.H.Identification of an abundant 56 kDa protein implicated in food allergy as granulebound starch synthase[J].Journal of Agricultural and Food Chemistry,2013,61(22):5 404-5 409.
[22]WANG X,FENG B,XU Z.B,et al.Identification and characterization of granule bound starch synthase I(GBSSI)gene of tartarybuckwheat(Fagopyrum tataricum Gaertn.)[J].Gene,2014,534(2):229-235.
[23]万峰,孙春玉,李闯,等.谷物胚乳中淀粉合成酶及其同工型的研究进展[J].分子植物育种,2017,15(11):126-135.WAN F,SUN C Y,LI C,et al.Research progress of starch synthase and its isoform in cereal endosperm[J].Molecular Plant Breeding,2017,15(11):126-135.
[24]FUNNELL-HARRIS D.L.,SATTLER S.E.,O’NEILL P.M.,et al.Effect of waxy(Low Amylose)on fungal infection of sorghum grain[J].Phytopathology,2015,105(6):786-796.
[25]熊飞,张琛,余徐润,等.专用小麦胚乳背部和腹部淀粉粒发育的差异[J].麦类作物学报,2013,33(6):1 270-1 276.XIONG F,ZHANG C,YU X R,et al.The differences in the development of starch granules in the back and abdomen of wheat endosperm[J].Journal of Triticeae Crops,2013,33(6):1 270-1 276.
[26]王忠,王慧慧,戴云霞,等.关于大麦胚乳细胞发育的研究[EB/OL].北京:中国科技论文在线,2012d.http://www.paper.edu.cn/releasepaper/content/201207-296.WANG Z,WANG H H,DAI Y X,et al.Study on the development of barley endosperm cells[EB/OL].Peking:Sciencepaper Online,2012d.http://www.paper.edu.cn/releasepaper/content/201207-296.
[27]侯维海,王建林,旦巴,等.西藏冬青稞胚乳淀粉粒发育特征研究[J].麦类作物学报,2017,37(9):1 232-1 239.HOU W H,WANG J L,DAN B,et al.Study on the development characteristics of starch granules of highland barley in Tibet[J].Journal of Triticeae Crops,2017,37(9):1 232-1239.
[28]罗伟,戴希碧.含水量对谷物热传导性能的影响[J].安徽农业科学,2010,38(21):11 453-11 454.LUO W,DAI X B.Influence of water content on the thermal conductivity performance of grain[J].Journal of Anhui Agricultural Sciences,2010,38(21):11 453-11 454.
[29]王剑平,盖玲,汪春,等.干谷物作为干燥介质的谷物吸收干燥过程平衡含水率模型[J].农业工程学报,1997,(2):220-224.WANG J P,GAI L,WANG J,et al.Equilibrium moisture content of adsorption grain drying using dried grain as adsorbents[J].Transactions of the Chinese Society of Agricultural Engineering,1997,(2):220-224.
[30]伊祖涛,张海艳.糯玉米胚乳发育过程中淀粉粒粒度分布的变化[J].江苏农业学报,2015,31(4):743-749.YI Z T,ZHANG H Y.Changes in the distribution of starch granule in the development of glutinous maize endosperm[J].Jiangsu Journal of Agricultural Sciences,2015,31(4):743-749.
[31]王兵,陈昕钰,徐德利,等.小麦胚乳充实与颖果运输组织的解剖学研究[J].山西农业大学学报(自然科学版),2017,37(12):841-848.WANG B,CHEN X Y,XU D L,et al.Study on the anatomy of wheat endosperm and caryopsis[J].Journal of Shanxi Agricultural University(Natural Science Edition),2017,37(12):841-848.
[2]杨智敏,孔德媛,杨晓云,等.青稞籽粒淀粉含量的差异[J].麦类作物学报,2013,33(6):1 139-1 143.YANG Z M,KONG D Y,YANG X Y,et al.The difference of starch content in highland barley[J].Journal of Triticeae Crops,2013,33(6):1 139-1 143.
[3]LI W H,XIAO X L,ZHANG W H,et al.Compositional,morphological,structural and physicochemical properties of starches from seven naked barley cultivars grown in China[J].Food Research International,2014,58(4):7-14.
[4]郑彦坤,曾德二,魏和平,等.水稻胚乳组织的结构观察[J].中国水稻科学,2017,31(1):91-98.ZHENG Y K,ZENG D E,WEI H P,et al.Structure observation of rice endosperm tissue[J].Chin.J.Rice Sci.,2017,31(1):91-98.
[5]王忠.水稻的开花与结实—水稻生殖器官发育图谱[M].北京:科学出版社,2015.
[6]李祥栋,石明,魏心元.禾谷类作物胚乳淀粉合成及Waxy基因研究进展[J].中国农学通报,2015,31(12):181-186.LI X D,SHI M,WEI X Y.Research progress of starch synthesis and Waxy gene in cereal crops[J].Chinese Agriculture Bulletin,2015,31(12):181-186.
[7]BERGER F.Endosperm development[J].Current Opinion in Plant Biology,1999,2(1):28-32.
[8]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):818-824.JIN Y P,LIU D T,LI D L,et al.The development of endosperm cells and their amyloid in wheat[J].Journal of Triticeae Crops,2013,33(4):818-824.
[9]VAN DONGEN J T,ROEB G W,DAUTZENBERG M,et al.Phloem import and storage metabolism are highly coordinated by the low oxygen concentrations within developing wheat seeds[J].Plant Physiology,2004,135(3):1 809-1 821.
[10]刘美,张凤,杨翠翠,等.小麦种子萌发早期淀粉降解关键酶活性及基因表达量研究[J].山东农业科学,2014,46(9):39-45.LIU M,ZHANG F,YANG C C,et al.Study on key enzyme activity and gene expression in early starch degradation of wheat seed germination[J].Shandong Agricultural Sciences,2014,46(9):39-45.
[11]SUH D S,VERHOEVEN T,DENYER K,et al.Characterization of Nubet and Franubet barley starches[J].Carbohydrate Polymers,2004,56(1):85-93.
[12]TANG H,MITSUNAGA T,KAWAMURA Y.Relationship between functionality and structure in barley starches[J].Carbohydrate Polymers,2004,57(2):145-152.
[13]高俊凤.植物生理学实验指导[M].北京:高等教育出版社,2006.
[14]张艺祎,李云霞,骆亮,等.低温储藏期间3种独蒜兰碳水化合物含量变化研究[EB/OL].北京:中国科技论文在线[2016-12-29].http://www.paper.edu.cn/releasepaper/content/201701-56.ZHANG Y Y,LI Y X,LUO L,et al.Study on the change of carbohydrate content of 3kinds of allicin in low temperature storage[EB/OL].PeKing:Sciencepaper Online[2016-12-29].http://www.paper.edu.cn/releasepaper/content/201701-56.
[15]余徐润,周亮,荆彦平,等.Image-Pro Plus软件在小麦淀粉粒显微图像分析中的应用[J].电子显微学报,2013,32(4):344-351.YU X R,ZHOU L,JING Y P,et al.Image-Pro Plus software used in wheat starch granule micrograph analysis[J].Journal of Chinese Electron Microscopy Society,2013,32(4):344-351.
[16]朱帆,徐广文,丁文平.表观直链淀粉含量对小麦淀粉凝胶流变特性影响的研究[J].食品科学,2008,29(2):43-47.ZHU F,XU G W,DING W P.Study on the effect of amylose content on the rheological properties of wheat starch gel[J].Food Science,2008,29(2):43-47.
[17]李栋梁,荆彦平,李小刚,等.高粱胚乳细胞与母体组织发育关系的研究[J].中国农业科学,2014,47(17):3 336-3 347.LI D L,JING Y P,LI X G,et al.Study on the relationship between sorghum endosperm cells and maternal tissue development[J].Scientia Agricultura Sinica,2014,47(17):3 336-3 347.
[18]王丹.夏橙果实成熟过程中果皮色泽变化的规律研究[D].武汉:华中农业大学,2016:1-2.
[19]荆彦平,李栋梁,刘大同,等.两个玉米品种胚乳发育的比较[J].玉米科学,2014,22(1):79-85.JING Y P,LI D L,LIU D T,et al.Comparison of endosperm development of two maize varieties[J].Journal of Maize Sciences,2014,22(1):79-85.
[20]苗红霞,孙佩光,张凯星,等.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[J].生物技术通报,2016,32(3):18-23.MIAO H X,SUN P G,ZHANG K X,et al.Research progress of the expression regulation mechanism of plant granule combined with starch synthase(GBSS)gene[J].Biotechnology Bulletin,2016,32(3):18-23.
[21]KRISHNAN H.B.,CHEN M.H.Identification of an abundant 56 kDa protein implicated in food allergy as granulebound starch synthase[J].Journal of Agricultural and Food Chemistry,2013,61(22):5 404-5 409.
[22]WANG X,FENG B,XU Z.B,et al.Identification and characterization of granule bound starch synthase I(GBSSI)gene of tartarybuckwheat(Fagopyrum tataricum Gaertn.)[J].Gene,2014,534(2):229-235.
[23]万峰,孙春玉,李闯,等.谷物胚乳中淀粉合成酶及其同工型的研究进展[J].分子植物育种,2017,15(11):126-135.WAN F,SUN C Y,LI C,et al.Research progress of starch synthase and its isoform in cereal endosperm[J].Molecular Plant Breeding,2017,15(11):126-135.
[24]FUNNELL-HARRIS D.L.,SATTLER S.E.,O’NEILL P.M.,et al.Effect of waxy(Low Amylose)on fungal infection of sorghum grain[J].Phytopathology,2015,105(6):786-796.
[25]熊飞,张琛,余徐润,等.专用小麦胚乳背部和腹部淀粉粒发育的差异[J].麦类作物学报,2013,33(6):1 270-1 276.XIONG F,ZHANG C,YU X R,et al.The differences in the development of starch granules in the back and abdomen of wheat endosperm[J].Journal of Triticeae Crops,2013,33(6):1 270-1 276.
[26]王忠,王慧慧,戴云霞,等.关于大麦胚乳细胞发育的研究[EB/OL].北京:中国科技论文在线,2012d.http://www.paper.edu.cn/releasepaper/content/201207-296.WANG Z,WANG H H,DAI Y X,et al.Study on the development of barley endosperm cells[EB/OL].Peking:Sciencepaper Online,2012d.http://www.paper.edu.cn/releasepaper/content/201207-296.
[27]侯维海,王建林,旦巴,等.西藏冬青稞胚乳淀粉粒发育特征研究[J].麦类作物学报,2017,37(9):1 232-1 239.HOU W H,WANG J L,DAN B,et al.Study on the development characteristics of starch granules of highland barley in Tibet[J].Journal of Triticeae Crops,2017,37(9):1 232-1239.
[28]罗伟,戴希碧.含水量对谷物热传导性能的影响[J].安徽农业科学,2010,38(21):11 453-11 454.LUO W,DAI X B.Influence of water content on the thermal conductivity performance of grain[J].Journal of Anhui Agricultural Sciences,2010,38(21):11 453-11 454.
[29]王剑平,盖玲,汪春,等.干谷物作为干燥介质的谷物吸收干燥过程平衡含水率模型[J].农业工程学报,1997,(2):220-224.WANG J P,GAI L,WANG J,et al.Equilibrium moisture content of adsorption grain drying using dried grain as adsorbents[J].Transactions of the Chinese Society of Agricultural Engineering,1997,(2):220-224.
[30]伊祖涛,张海艳.糯玉米胚乳发育过程中淀粉粒粒度分布的变化[J].江苏农业学报,2015,31(4):743-749.YI Z T,ZHANG H Y.Changes in the distribution of starch granule in the development of glutinous maize endosperm[J].Jiangsu Journal of Agricultural Sciences,2015,31(4):743-749.
[31]王兵,陈昕钰,徐德利,等.小麦胚乳充实与颖果运输组织的解剖学研究[J].山西农业大学学报(自然科学版),2017,37(12):841-848.WANG B,CHEN X Y,XU D L,et al.Study on the anatomy of wheat endosperm and caryopsis[J].Journal of Shanxi Agricultural University(Natural Science Edition),2017,37(12):841-848.