外源ABA和6-BA对不同持绿型小麦籽粒灌浆进程及蛋白质含量影响

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杨东清, 李玉玲, 倪英丽, 尹燕枰, 杨卫兵, 崔正勇, 张永太, 马仁元, 王振林. 外源ABA和6-BA对不同持绿型小麦籽粒灌浆进程及蛋白质含量影响. 作物学报, 2014, 40(2): 301-312[YANG Dong-Qing, LI Yu-Ling, NI Ying-Li, YIN Yan-Ping, YANG Wei-Bing, CUI Zheng-Yong, ZHANG Yong-Tai, MA Ren-Yuan, WANG Zhen-Lin. Effects of Exogenous ABA and 6-BA on Protein Content and Grain Filling Process in Different Types of Stay-Green Wheat. 作物学报, 2014, 40(2): 301-312] 复制到剪切板

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外源ABA和6-BA对不同持绿型小麦籽粒灌浆进程及蛋白质含量影响

杨东清¹, 李玉玲², 倪英丽^1,³, 尹燕枰¹, 杨卫兵¹, 崔正勇¹, 张永太², 马仁元², 王振林^1,^*

¹山东农业大学农学院 / 作物生物学国家重点实验室, 山东泰安 271018

²齐河县农业局, 山东齐河 251100

³任城区农业局, 山东济宁 272000

^* 通讯作者(Corresponding author): 王振林, E-mail:zlwang@sdau.edu.cn, Tel: 0538-8241359

收稿日期:2013-09-06

基金:本研究由国家公益性行业(农业)科研专项经费项目(201203100, 201203029), 国家自然科学基金项目(31271661), 国家重点基础研究发展计划(973计划)项目(2009CB118602), 国家粮食丰产科技工程项目(2012BAD04B05)和山东省现代农业技术体系项目经费资助。

摘要

本文旨在揭示持绿型小麦品种的灌浆特性和籽粒蛋白质含量对外源激素的响应, 认识植物激素调控籽粒灌浆和蛋白质合成的作用机制。自盛花期开始, 连续4 d喷施10 mg L^-1脱落酸(ABA)和6-苄基腺嘌呤(6-BA), 持绿型品种汶农6号的灌浆期(t₃)、活跃生长期(D)、平均灌浆速率(G_mean)、最大灌浆速率(G_max)、千粒重(TGW)和产量(GY)均大于非持绿型济麦20, 且强势粒的各项灌浆参数均大于弱势粒。喷施两种外源激素显著提高两品种的G_mean、G_max、TGW和GY, 对t₃和D的影响存在粒位及品种效应。喷施ABA和6-BA均显著提高花后35 d的籽粒蛋白质含量, 显著提高汶农6号强、弱势粒的麦谷蛋白含量, 但济麦20强、弱势粒的麦谷蛋白含量显著降低。喷施ABA后, 汶农6号强势粒花后7~21 d的玉米素核苷(ZR)含量显著升高; 喷施6-BA后, 济麦20弱势粒花后14~35 d的ZR含量显著升高, 而汶农6号强势粒7~21 d的赤霉素(GA₃)含量显著降低, 弱势粒花后7~28 d的GA₃含量显著提高。喷施外源ABA和6-BA显著提高两品种强势粒花后7~21 d的生长素(IAA)和ABA含量。ABA和6-BA处理显著提高花后7~21 d谷氨酰胺合成酶(GS)活性。可见, 喷施外源ABA和6-BA使小麦内源激素水平变化, 促进籽粒灌浆, 增加粒重和产量, 提高籽粒GS活性和蛋白质含量, 改变蛋白质组分。

关键词: 持绿型小麦; 激素; 灌浆特性; Richard方程; 蛋白质含量; 谷氨酰胺合成酶

Effects of Exogenous ABA and 6-BA on Protein Content and Grain Filling Process in Different Types of Stay-Green Wheat

YANG Dong-Qing¹, LI Yu-Ling², NI Ying-Li^1,³, YIN Yan-Ping¹, YANG Wei-Bing¹, CUI Zheng-Yong¹, ZHANG Yong-Tai², MA Ren-Yuan², WANG Zhen-Lin^1,^*

¹ College of Agronomy, Shandong Agricultural University / State Key Laboratory of Crop Biology, Tai#cod#x02019;an 271018, China

² Agriculture Bureau of Qihe County of Shandong Province, Qihe 251100, China

³ Agriculture Bureau of Rencheng District of Shandong Province, Jining 272000, China

Abstract

This study aimed to understand the grain filling process and protein biosynthesis in grain of stay-green wheat in response to exogenous hormones. At blooming stage, plants of Wennong 6 (stay-green cultivar) and Jimai 20 (control) were sprayed with 10 mg L^-1 abscisic acid (ABA) and 6-benzyladenine (6-BA) for four days. Compared with Jimai 20, Wennong 6 had longer grain-filling duration (t₃) and the active growth phase (D), higher mean grain-filling rate (G_mean) and maximum grain-filling rate (G_max), larger thousand-grain weight (TGW), and greater grain yield (GY). All parameters for grain filling were higher in superior grain than in inferior grain.G_mean,G_max, TGW, and GY were promoted by ABA or 6-BA treatment in both cultivars. However, the effects of exogenous hormones ont₃ andD were interacted with grain position in a panicle and genotype. Exogenous ABA and 6-BA treatment obviously increased protein content at 35 days after anthesis (DAA). The glutenin contents in ABA-treated and 6-BA-treated Wennong 6 significantly increased at 35 DAA; however, those in Jimai 20 decreased at 35 DAA. Exogenous ABA increased endogenous zeatin (ZR) content from 7 DAA to 21 DAA. In Wennong 6, GA₃ content in superior grain was significantly decreased from 7 DAA to 21 DAA by 6-BA treatment, however, its content in inferior grain was enhanced. In both cultivars, application of ABA and 6-BA resulted in significant increases of endogenous auxin (IAA) and ABA contents from 7 DAA to 21 DAA, along with significant enhancement of glutamine synthetase (GS) activity.

Keyword: Stay-green wheat; Hormones; Grain filling characteristics; Richard equation; Protein content; Glutamine synthetase

Show Figures

小麦籽粒中蛋白质的含量和组分对小麦的产量、营养品质及加工品质均有重要影响^{[ 1]}。研究小麦籽粒灌浆进程中蛋白质含量动态变化以及蛋白质组分对于小麦品质有重要意义。植物激素是植物自身合成的生长调节物质, 对植物的生长发育发挥重要的调控作用^{[ 2]}。当施用植物激素或植物生长调节剂后, 作物的氮代谢受到调控。烯效唑拌种提高了小麦叶片全氮含量^{[ 3]}; 细胞分裂素可诱导与蛋白质合成有关基因的表达, 调节蛋白质的合成^{[ 4]}; 小麦籽粒中玉米素核苷(ZR)含量与蛋白质含量呈显著正相关^{[ 5]}; ZR在盛花期能促进棉花对氮素的吸收, 而降低结铃期后棉株ABA含量有利于氮素的吸收利用^{[ 6]}; GA₃可降低大豆根系硝态氮含量, 对硝酸还原酶有一定抑制作用^{[ 7]}; GA₃对棉铃Bt蛋白含量的调节效应与棉铃所处发育阶段有关^{[ 8]}; 外源ABA可增加籽粒中蛋白质含量, 改变谷蛋白组分含量^{[ 9]}。

研究表明, 持绿型作物叶片叶绿素含量高, 植株衰老过程中叶片失绿缓慢^{[ 10]}, 选育保绿品种是提高作物产量的有效途径^{[ 11, 12]}。持绿型小麦绿叶面积大, 叶片光合能力强, 有较长的灌浆持续期^{[ 13]}。不同持绿型小麦灌浆进程以及产量形成的生物学机制有显著差异^{[ 14, 15]}。然而, 关于不同持绿型小麦品种籽粒灌浆进程及其蛋白质含量动态以及激素调控机制还有待深入研究。本试验选用不同持绿型小麦品种进行花后喷施外源ABA和6-BA, 并对照非持绿型品种分析籽粒灌浆进程、蛋白质含量、谷氨酰胺合成酶活性、成熟期籽粒蛋白质组分含量以及内源激素含量变化, 旨在研究持绿型小麦品种灌浆进程中蛋白质含量和组分及其合成关键酶的变化特性, 为以化学调控措施实现小麦高产、优质生产提供理论依据。

1 材料与方法

1.1 试验设计

2010—2011和2011—2012小麦生长季, 在山东农业大学农学试验农场(山东泰安)进行田间试验。试验地耕层(0~20 cm)土壤含有机质12.3 g kg^-1、全氮0.91 g kg^-1、碱解氮87.2 mg kg^-1、速效磷8.6 mg kg^-1、速效钾57.5 mg kg^-1。播种前施入基肥纯氮120 kg hm^-2、P₂O₅100 kg hm^-2、K₂O 100 kg hm^-2, 拔节期追施纯氮120 kg hm^-2。其他田间管理同一般高产田。试验小区面积9 m², 随机排列, 重复3次。

采用二因素随机区组设计。以品种为主区, 选用持绿型汶农6号和非持绿型济麦20; 花后喷施外源激素为副区, 设ABA、6-BA和清水(CK) 3个处理。于盛花期开始, 每天16:00左右在穗部喷施10 mg L^-1 ABA或6-BA, 每个处理喷施9 m², 用量为100 mL m^-2, 连续喷4 d。为使激素能更好地附着于穗部, 激素溶液中加入0.5% Tween-20, CK为含Tween-20清水。

于开花期标记长相一致、同一日开花的麦穗。开花后7、14、21、28和35 d分别取中部小穗的第1、第2位籽粒(强势粒)和第3、第4位籽粒(弱势粒)。若无第4位籽粒, 则只取第3位籽粒为弱势粒; 若有第5位籽粒, 则取第3、第4、第5位籽粒为弱势粒。一半样品保存于-40℃冰箱中, 用于测定谷氨酰胺合成酶活性、内源激素含量测定; 另一半于55℃烘干至恒重, 用于测定蛋白质含量以及蛋白质组分。花后3、6、9、12、15、18、21、24、28和35 d (弱势粒从花后6 d开始)取穗样, 烘干后分强、弱势粒作籽粒灌浆动态模拟。

小麦成熟后每处理收获1 m²统计穗数, 脱粒后晾干称重, 计算产量。3次重复。

1.2 籽粒增重及灌浆参数的计算

按朱庆森等^{[ 16]}介绍的方法, 利用Richard方程模拟小麦籽粒增重动态。

(1)

式中, W为粒重, A为生长终止量, t为开花后天数, B、 k、 N为参数。对该方程求一阶导数, 可得灌浆速率( G)。

(2)

当灌浆进程达到 A的99%时, 实际灌浆终期( t₃)、最大灌浆速率( G_max)、平均灌浆速率( G_mean)和活跃生长期( D)分别用下列公式计算:

(3)

(4)

(5)

(6)

1.3 测定项目与方法

采用半微量凯氏定氮法测定氮含量, 再乘以系数5.7, 即为小麦籽粒粗蛋白质含量^{[ 17]}。参照何照范^{[ 17]}的方法分离和测定蛋白质组分。按王平等^{[ 18]}的方法测定谷氨酰胺合成酶(GS)活性。

采用高效液相色谱法(HPLC)测定籽粒中4种内源激素的含量^{[ 19]}。以体积比为50∶45∶5的甲醇∶乙腈∶0.6%乙酸作为流动相; 分析柱为Symmetry C18 (150.0 mm × 4.6 mm, 5 μm), 柱温为25℃, 流动相流速0.6 mL min^-1; 检测波长为254 nm。玉米素、赤霉素、生长素、脱落酸标样均购自美国Sigma公司。

1.4 统计分析

小麦籽粒产量为2年度数据, 而籽粒灌浆进程、蛋白质含量及组分、GS活性、内源激素含量均为2010—2011年度数据。用DPS7.05统计分析软件对数据进行方差分析和显著性检验以及相关性分析, 用MATLAB 7.0计算籽粒灌浆参数, 用Sigma 10.0作图。

2 结果与分析

2.1 外源ABA和6-BA对籽粒灌浆动态及灌浆速率的影响

两品种强、弱势籽粒增重过程均呈S曲线变化, 花后35 d粒重达到最大, 但此时持绿型品种的粒重高于非持绿型品种(图1)。喷施外源ABA和6-BA提高了两品种强、弱势粒粒重, ABA处理汶农6号花后35 d的强、弱势粒粒重分别提高5.93%和5.84%, 低于济麦20的15.01%和14.44%; 而6-BA处理后强、弱势粒粒重增幅在两品种间差异较小, 汶农6号的增幅分别为6.88%和6.55%, 济麦20的增幅分别为7.06%和9.11%。

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图1 外源ABA和6-BA对籽粒增重的影响实心图例表示强势粒(S), 空心图例表示弱势粒(I), 虚线表示对照; CK、ABA和6-BA分别表示喷施清水(含Tween-20)、ABA (10 mg L^-1)和6-BA (10 mg L^-1)。Fig. 1 Effects of spraying exogenous ABA and 6-BA on grain weight processSolid and hollow legends represent superior (S) and inferior (I) grain, respectively. Dash lines represent the control (CK). CK, ABA, and 6-BA represent spraying water (including Tween-20), ABA (10 mg L^-1), and 6-BA (10 mg L^-1), respectively.

两品种籽粒灌浆速率呈先升高再降低的趋势。济麦20灌浆速率在花后15 d达最大, 而汶农6号则在花后18 d达最大(图2)。喷施外源ABA, 汶农6号强势粒花后6~15 d的灌浆速率降低, 但强势粒花后18~28 d和弱势粒花后6~28 d的灌浆速率提高; 而济麦20强、弱势粒的灌浆速率均在花后6~35 d较CK提高。6-BA处理提高了汶农6号强势粒花后3~21 d和弱势粒6~15 d的灌浆速率, 对济麦20强势粒的作用时段与汶农6号相同, 而对弱势粒的作用时间更长(花后6~35 d)。

2.2 外源ABA和6-BA对小麦籽粒灌浆参数的影响

持绿型汶农6号强、弱势籽粒的 t₃、 D、 G_max和 G_mean参数值均大于非持绿型济麦20 (表1)。两品种强势粒的各灌浆参数均大于弱势粒。ABA处理后, 汶农6号强势粒的 t₃和 D值没有明显变化, 但强、弱势粒的 G_max和 G_mean明显升高; 6-BA处理后, 汶农6号的 t₃和 D呈现强势粒减小, 而弱势粒增大的变化特点。与汶农6号不同的是, 两种外源激素处理后济麦20强、弱势粒的上述参数均呈增大趋势。这些结果表明ABA和6-BA对灌浆参数的影响存在粒位和品种效应。

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图2 外源ABA和6-BA对籽粒灌浆速率的影响实心图例表示强势粒(S), 空心图例表示弱势粒(I), 虚线表示对照; CK、ABA和6-BA分别表示喷施清水(含Tween-20)、ABA (10 mg L^-1)和6-BA (10 mg L^-1)。Fig. 2 Effects of spraying exogenous ABA and 6-BA on grain filling rateSolid and hollow legends represent superior (S) and inferior (I) grain, respectively. Dash lines represent the control (CK). CK, ABA, and 6-BA represent spraying water (including Tween-20), ABA (10 mg L^-1), and 6-BA (10 mg L^-1), respectively.

表1 籽粒增重模型与籽粒灌浆参数 Table 1 The grain filling parameters and grain filling process model

2.3 不同激素处理对小麦产量的影响

对照条件下, 汶农6号的千粒重和产量均显著大于济麦20, 表明持绿型品种产量较高。喷施ABA和6-BA显著提高两品种的千粒重和产量, 尤其是ABA处理, 汶农6号增产幅度大于济麦20。2010—2011年度ABA和6-BA处理的增产幅度, 汶农6号分别为14.1%和6.2%, 济麦20分别为5.3%和3.1%。两种外源激素处理对穗数和穗粒数没有显著影响(表2)。

2.4 外源ABA和6-BA对籽粒蛋白质含量的影响

两品种强弱势籽粒蛋白质含量变化趋势相同, 均先降低后升高(图3)。济麦20的蛋白质含量高于汶农6号。花后35 d, 两品种强势粒蛋白质含量均高于弱势粒。喷施ABA和6-BA显著提高花后35 d的籽粒蛋白质含量, 且两品种强、弱势粒的蛋白质含量与对照的差异缩小。表明采用外源激素可以改善籽粒蛋白质含量。

表2 激素处理对不同品种小麦产量和产量构成的影响 Table 2 Effects of exogenous hormones treatments on the factors of wheat yield

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	图3 外源ABA与6-BA对强势粒(A、C)和弱势粒(B、D)蛋白质含量的影响Fig. 3 Effect of ABA and 6-BA on protein content in superior (A, C) and inferior (B, D) grain

2.5 喷施外源ABA和6-BA对籽粒蛋白质组分含量的影响

小麦籽粒中各蛋白质组分所占比例不同, 以麦谷蛋白含量最高。非持绿型济麦20强势粒的清蛋白、球蛋白、醇溶蛋白显著高于持绿型汶农6号。

在对照条件下, 汶农6号强势粒的清蛋白、醇溶蛋白、麦谷蛋白含量显著高于弱势粒; 而济麦20强势粒的清蛋白、球蛋白显著低于弱势粒, 醇溶蛋白、麦谷蛋白含量显著高于弱势粒。喷施ABA显著提高了两品种强势粒的清蛋白、球蛋白、醇溶蛋白含量; 6-BA处理显著降低汶农6号强、弱势粒清蛋白含量, 但显著提高济麦20强、弱势粒的清蛋白含量。ABA和6-BA处理后, 麦谷蛋白含量汶农6号显著升高, 而济麦20显著降低, 且强、弱势粒表现一致(图4)。可见, 激素处理对蛋白质组分含量的影响存在品种和粒位效应。

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图4 外源ABA和6-BA对小麦籽粒蛋白质组分含量的影响S表示强势粒, I表示弱势粒; CK、ABA和6-BA分别表示喷施清水(含Tween-20)、ABA (10 mg L^-1)和6-BA (10 mg L^-1)。Fig. 4 Effect of ABA and 6-BA on protein components in wheatS and I represent superior and inferior grain, respectively. CK, ABA, and 6-BA represent spraying water (including Tween-20), ABA (10 mg L^-1), and 6-BA (10 mg L^-1), respectively.

2.6 内源激素含量动态

2.6.1 玉米素核苷(ZR) 两品种在灌浆期强、弱势籽粒的ZR含量均呈先升高后降低的趋势, 花后14 d达最高值(图5-A, B)。与对照相比, ABA处理后济麦20强势粒的ZR含量, 前期(14~21 d)降低, 后期(28~35 d)升高, 而弱势粒的ZR含量则持续(7~ 35 d)升高; 汶农6号强势粒ZR含量升高期为花后7~21 d, 比济麦20提早7 d。喷施6-BA显著提高济麦20弱势籽粒花后14~35 d的ZR含量, 且显著提高汶农6号强、弱势籽粒花后14~28 d的ZR含量。

2.6.2 赤霉素(GA₃) 两品种籽粒GA₃含量呈先升高后降低的变化趋势, 但强、弱势粒的峰值时间不同, 强势粒为花后14 d, 弱势粒为花后21 d(图5-C, D)。外源ABA处理对汶农6号强势粒花后7 d的GA₃含量无显著影响, 但显著降低花后14 d的GA₃含量; 显著提高汶农6号强、弱势粒花后21~35 d的GA₃含量。6-BA处理显著降低汶农6号强势粒7~21 d的GA₃含量, 但显著提高弱势粒花后7~28 d的GA₃含量。

2.6.3 生长素(IAA) 小麦籽粒中IAA含量随灌浆进程呈先升高后降低趋势, 两品种强、弱势粒分别在花后21 d和14 d达峰值, 强势粒的IAA含量显著高于弱势粒。外源ABA和6-BA显著提高两品种强势粒的IAA含量, 作用时间不同, 前者为花后7~35 d, 后者为花后7~21 d; 而弱势粒的IAA含量在花后7~35 d持续增大, 两品种表现一致(图5-E, F)。

2.6.4 脱落酸(ABA) 与其他籽粒内源激素表现不同, ABA含量在灌溉过程中出现两次升高-降低的变化, 强、弱势粒峰值时间一致。ABA最大峰值出现在花后14 d, 第二峰值在花后28 d, 而花后21 d和35 d达到两个低谷值。花后7~14 d, 济麦20强势粒的ABA含量高于弱势粒, 但花后21~35 d, 弱势粒的ABA含量高于强势粒。花后7~35 d, 汶农6号弱势粒ABA含量显著高于强势粒。喷施ABA显著提高持绿型汶农6号花后7~28 d强、弱势籽粒及济麦20弱势粒中ABA含量。喷施6-BA显著提高汶农6号弱势粒花后7~28 d的ABA含量, 显著提高济麦弱势粒7~21 d的ABA含量(图5-G, H)。

2.7 外源ABA和6-BA对籽粒GS活性的影响

所有处理中GS活性均呈先升高后降低的变化趋势, 花后14 d达最大值; 济麦20的GS活性高于汶农6号(图8)。两品种强、弱势籽粒的GS活性存在显著差异, 花后7~21 d强势粒的GS活性高于弱势粒。ABA和6-BA均显著提高花后7~14 d的GS活性, 在花后28~35 d没有显著效应。ABA处理后, 强势粒花后7~14 d的GS活性相对较高, 6-BA处理花后28 d的GS活性下降相对缓慢, 保持较高的活性。

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图5 不同外源激素对小麦叶片内源激素含量的影响实心图例表示强势粒(S), 空心图例表示弱势粒(I), 虚线表示对照; CK、ABA和6-BA分别表示喷施清水(含Tween-20)、ABA (10 mg L^-1)和6-BA (10 mg L^-1)。Fig. 5 Effects of exogenous hormones on content of four endogenous hormones in wheat grainsSolid and hollow legends represent superior (S) and inferior (I) grain, respectively. Dash lines represent the control (CK). CK, ABA, and 6-BA represent spraying water (including Tween-20), ABA (10 mg L^-1), and 6-BA (10 mg L^-1), respectively.

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	图6 外源ABA和6-BA对强势粒(A、C)和弱势粒(B、D)中GS活性的影响Fig. 6 GS activity in response to exogenous ABA and 6-BA in superior (A, C) and inferior (B, D) grain

3 讨论

持绿型汶农6号的灌浆终期及活跃生长期均大于非持绿品种济麦20, 且汶农6号最大灌浆速率及平均灌浆速率均大于济麦20, 因而汶农6号在较长的灌浆期内能积累更多的同化物, 最终获得较高的粒重及产量(表2)。这进一步证实持绿型小麦品种由于叶片功能期较长, 籽粒灌浆时间长, 为籽粒灌浆提供充足的营养物质, 产量一般高于非持绿品种^{[ 15, 20, 21]}。

植物激素在作物生长发育中起着重要的调控作用, 同化物质的转运及在库中的积累转化都有激素参与调控^{[ 22, 23]}。外源ABA和细胞分裂素参与调控籽粒灌浆及籽粒中的物质代谢^{[ 24, 25]}, ABA参与光合产物的合成和积累^{[ 26]}。研究表明, ABA促进氮素向籽粒的转运, 提高籽粒的蛋白质含量^{[ 27]}。ABA促进贮藏物质的积累, 特别是贮藏蛋白的合成^{[ 28, 29]}, 贮藏蛋白的基因受ABA信号的诱导而提高表达水平^{[ 30]}。也有研究认为ABA对贮藏蛋白合成的促进作用发生在转录水平上^{[ 31]}。本试验中喷施外源ABA显著提高了两品种籽粒花后35 d的蛋白质含量。谷氨酰胺合成酶是合成蛋白质的关键酶之一, 喷施ABA提高了GS活性(图6), 因而有利蛋白质的合成, 这与Yang等^{[ 32]}的报道一致, 即ABA通过提高GS活性促进蛋白质的合成。6-BA能提高小麦花后同化氮素对籽粒总氮的贡献率^{[ 33]}。Criado等^{[ 34]}发现, 6-BA能够抑制叶片蛋白质的降解及促进蛋白质的合成。Ananieva等^{[ 35]}研究表明, 6-BA可促进茭瓜子叶中蛋白质和RNA的合成。在本研究中, 外源6-BA提高籽粒花后7~14 d的GS活性, 且显著提高花后35 d蛋白质含量。这表明外源ABA和6-BA均能提高GS活性, 提高小麦籽粒蛋白质含量。蛋白质的组分分为清蛋白、球蛋白、醇溶蛋白和麦谷蛋白^{[ 36]}。研究表明, 6-BA处理小麦穗可提高小麦谷蛋白/醇溶蛋白比值^{[ 37]}。本研究发现, 喷施ABA显著提高两品种强势粒清蛋白、球蛋白、醇溶蛋白含量; 6-BA处理显著降低汶农6号强、弱势粒清蛋白含量, 但显著提高济麦20强、弱势粒清蛋白含量。此外, 也证实ABA处理显著提高汶农6号强、弱势籽粒的麦谷蛋白含量^{[ 9]}。

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	图7 HPLC法测定4种激素的标样图Fig. 7 The standard records of four hormones determined by HPLC

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	图8 HPLC法测定籽粒中4种内源激素的记录图Fig. 8 Records of four endogenous hormones in grains determined by HPLC

籽粒灌浆过程中各内源激素的含量在品种间存在基因型差异^{[ 38]}。外源激素通过改变内源激素水平调节植物内在生理代谢的各个环节^{[ 39]}。本试验中喷施ABA和6-BA不同程度地改变了两品种内源ZR、ABA、GA₃、IAA含量, 且存在明显的品种和粒位效应。植物激素含量变化由基因表达调控^{[ 40]}, 作物的持绿特性由细胞核及细胞质基因控制^{[ 41]}。持绿型汶农6号和非持绿济麦20的内源激素对ABA和6-BA的响应不同, 这可能是由两品种遗传基因的差异性造成的。研究表明, GA的生物合成受GA含量的反馈调节, 低含量的GA会激活GA的生物合成^{[ 42]}。外源ABA能提高内源ABA的含量^{[ 19]}, 这一点也在本试验的两个品种中得到验证。ABA的生物合成类萜途径是由甲瓦龙酸(MVA)经中间产物异戊烯基焦磷酸(IPP)等一系列步骤^{[ 43, 44]}, 而IPP也是玉米素生物合成的起始物^{[ 45]}, 因而本试验中喷施ABA提高ZR含量可能是通过促使IPP向玉米素合成实现的, 而外源ABA提高GA₃含量可能是通过降低前期内源GA₃含量, 由GA₃的反馈调节提高籽粒灌浆后期的GA₃含量实现的。

内源细胞分裂素可以提高籽粒灌浆强度, 增大千粒重^{[ 46]}。细胞分裂素与其特定受体结合后诱导基因表达, 促进RNA和蛋白质的合成^{[ 47]}。GA对籽粒灌浆起调控作用, 且与粒重的决定密切相关^{[ 25]}。也有研究认为, GA可诱导产生或激活α-淀粉酶等水解酶而不利于淀粉和蛋白质的合成和积累^{[ 48]}。ABA表现多种生理功能, 如促进水稻籽粒灌浆^{[ 49]}; 延长小麦叶片保绿时间, 延缓叶绿素降解^{[ 50]}; 提高小麦穗颈节伤流强度, 促进营养物质的运转^{[ 51]}; 抑制一些水解酶的活性, 并促进蛋白质的合成和积累^{[ 25]}。IAA对胚乳细胞的发育以及籽粒中物质积累有调控作用^{[ 52]}。外源ABA和6-BA对籽粒灌浆进程和蛋白质含量影响的机制可能在于, 外源激素处理引起籽粒中激素含量和比例变化, 导致蛋白质合成与分解的关键酶活性改变。本研究中, 喷施外源ABA和6-BA提高了两品种籽粒平均及最大灌浆速率, 但对灌浆期和活跃生长期的影响存在品种差异, ABA处理后, 汶农6号强势粒的灌浆期及活跃生长期没有明显变化。这可能是非持绿型品种叶片叶绿素含量低, 抗氧化性弱, 衰老快, 导致其功能期较短, 因而ABA和6-BA能延长非持绿型济麦20的灌浆期。

4 结论

持绿型汶农6号的 t₃、 D、 G_mean和 G_max均大于非持绿型济麦20, 使汶农6号籽粒能积累更多的光合产物, 因而汶农6号粒重及产量较大。喷施外源ABA和6-BA使籽粒内源激素不同程度地改变, 平均灌浆速率和最大灌浆速率提高, 促进籽粒灌浆、千粒重和产量提高。ABA和6-BA处理后, GS活性及蛋白质含量均提高, 蛋白质组分较对照有不同程度改变。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。The authors have declared that no competing interests exist.

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25	杨建昌, 王志琴, 朱庆森, 苏宝林. ABA与GA对水稻籽粒灌浆的调控. 作物学报, 1999, 25: 341-348 Yang J C, Wang Z Q, Zhu Q S, Su B L. Regulation of ABA and GA to the grain filling of rice. Acta Agron Sin, 1999, 25: 341-348 (in Chinese with English abstract) [本文引用:3] [CJCR: 1.667]
26	刘晓冰, 金剑, 王光华, Herbert S J. 调节大豆粒重的生理生态因素. 大豆科学, 2007, 26: 607-617 Liu X B, Jin J, Wang G H, Herbert S J. Regulation of soybean seed size by eco-physiological factors. Soybean Sci, 2007, 26: 607-617 (in Chinese with English abstract) [本文引用:1]
27	Xie Z J, Jiang D, Dai T B, Jing Q, Cao W X. Effects of exogenous ABA and cytokinin on leaf photosynthesis and grain protein accumulation in wheat ears cultured in vitro. Plant Growth Regul, 2004, 44: 25-32 [本文引用:1] [JCR: 1.67]
28	宋松泉, 段永新, 傅家瑞. ABA对种子发育的调节. 种子, 1997, (5): 36-42 Song S Q, Duan Y X, Fu J R. Regulation of ABA on seed development. Seed, 1997, (5): 36-42 (in Chinese with English abstract) [本文引用:1] [CJCR: 0.445]
29	Finkelstein R R, Gibson S I. ABA and sugar interactions regulating development: cross-talkor voices in a crowd. Curr Opin Plant Biol, 2002, 5: 26-32 [本文引用:1] [JCR: 8.455]
30	李杉, 邢更妹, 崔凯荣, 王亚馥. 植物体细胞胚发生中ATP酶活性时空分布动态与内源激素的变化. 植物学通报, 2001, 18: 308-317 Li S, Xing G M, Cui K R, Wang Y F. ATPase activity and endogenous hormone changes during plant somatic embryogenesis. Chin Bull Bot, 2001, 18: 308-317 (in Chinese with English abstract) [本文引用:1]
31	黄上志, 傅家瑞. 脱落酸对发育中花生胚萌发和贮藏蛋白质合成的影响. 植物生理学报, 1993, 19: 31-37 Huang S Z, Fu J R. Effect of abscisic acid on precocious germination and storage protein during culture of developing peanut embryos. Acta Phytophysiol Sin, 1993, 19: 31-37 (in Chinese with English abstract) [本文引用:1]
32	Yang W B, Cai T, Ni Y L, Li Y, Guo J X, Peng D L, Yang D Q, Yin Y P, Wang Z L. Effects of exogenous abscisic acid and gibberellic acid on filling process and nitrogen metabolism characteristics in wheat grains. Aust J Crop Sci, 2013, 7: 58-65 [本文引用:1] [JCR: 1.632]
33	谢祝捷, 姜东, 曹卫星, 戴廷波, 荆奇. 花后干旱和渍水条件下生长调节物质对冬小麦光合特性和物质运转的影响. 作物学报, 2004, 30: 1047-1052 Xie Z J, Jiang D, Cao W X, Dai T B, Jing Q. Effects of plant growth regulation substances on photosynthetic characteristics and assimilate transportation in winter wheat under post-anthesis drought and waterlogging. Acta Agron Sin, 2004, 30: 1047-1052 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.667]
34	Criado M V, Caputo C, Roberts I N, Castro M A, Barneix A J. Cytokinin-induced changes of nitrogen remobilization and chloroplast ultrastructurein wheat (Triticum aestivum). J Plant Physiol, 2009, 166: 1775-1785 [本文引用:1] [JCR: 2.699]
35	Ananieva K I, Ananiev E D. Effect of methyl ester of jasmonic acid and benzylaminopurine on growth and protein profile of excised cotyledons of Cucurvita pepo (Zucchini). Biol Plant, 1999, 42: 549-557 [本文引用:1] [JCR: 2.32]
36	高文川, 马猛, 王爱娜, 赵慧贤. 不同品质类型小麦籽粒麦谷蛋白亚基及谷蛋白聚合体形成和累积动态. 作物学报, 2010, 36: 1769-1776 Gao W C, Ma M, Wang A N, Zhao H X. Accumulation dynamics of glutenin suunits and glutenin polymer in wheat cultivars with different qualities during grain development. Acta Agron Sin, 2010, 36: 1769-1776 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.667]
37	张敏, 戴廷波, 姜东, 荆奇, 曹卫星. 6-BA对小麦花后C/N物质运转和籽粒品质的影响. 南京农业大学学报, 2006, 29(4): 6-10 Zhang M, Dai T B, Jiang D, Jing Q, Cao W X. Effects of 6-BA on carbon and nitrogen assimilate translocation after anthesis and quality formation in wheat. J Nanjing Agric Univ, 2006, 29(4): 6-10 (in Chinese with English abstract) [本文引用:1] [CJCR: 0.916]
38	李文阳, 尹燕枰, 严素辉, 戴忠民, 梁太波, 王振林. 不同粒型小麦品种籽粒内源激素变化与籽粒灌浆特征的比较. 华北农学报, 2007, 22(1): 5-8 Li W Y, Yin Y P, Yan S H, Dai Z M, Liang T B, Wang Z L. Hormonal changes in relation to filling characters in developing grains of two wheat cultivars differing in grain size. Acta Agric Boreali-Sin, 2007, 22(1): 5-8 (in Chinese with English abstract) [本文引用:1] [CJCR: 0.951]
39	郑莎莎, 孙传范, 孙红春, 刘连涛, 赵金峰, 李存东. 不同外源激素对花铃期棉花主茎叶生理特性的影响. 中国农业科学, 2009, 42: 4383-4389 Zheng S S, Sun C F, Sun H C, Liu L T, Zhao J F, Li C D. Effects of different exogenous hormones on physiological characteristics of main stem leaves at flower and boll stage in cotton. Sci Agric Sin, 2009, 42: 4383-4389 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.889]
40	Wilkinson S, Kudoyarova G R, Veselov D S, Arkhipova T N, Davies W J. Plant hormone interactions: innovative targets for crop breeding and management. J Exp Bot, 2012, 63: 3499-3509 [本文引用:1] [JCR: 5.242]
41	Cha K W, Lee Y J, Koh H J, Lee B M, Nam Y W, Paek N C. Isolation, characterization, and mapping of the stay green mutant in rice. Theor Appl Genet, 2002, 104: 526-532 [本文引用:1] [JCR: 3.658]
42	Yamaguchi S, Kamiya Y J. Gibberellin biosynthesis: its regulation by endogenous and environmental signals. Plant Cell Physiol, 2000, 41: 251-257 [本文引用:1] [JCR: 4.134]
43	Cutler A J, Krochko J E. Formation and breakdown of ABA. Trends Plant Sci, 1999, 4: 472-478 [本文引用:1] [JCR: 11.808]
44	Seo M, Koshiba T. Complex regulation of ABA biosynthesis in plants. Trends Plant Sci, 2002, 7: 41-48 [本文引用:1] [JCR: 11.808]
45	王忠. 植物生理学. 北京: 中国农业出版社, 2006. pp282-283 Wang Z. Plant Physiology. Beijing: China Agriculture Press, 2006. pp282-283(in Chinese) [本文引用:1]
46	李秀菊, 职明星, 鲁玉贞. 不同粒型小麦籽粒发育过程中细胞分裂素含量的变化. 麦类作物学报, 2005, 25(2): 42-45 Li X J, Zhi M X, Lu Y Z. Changes of cytokinins content during the wheat grain development of different grain sizes. J Triticeae Crops, 2005, 25(2): 42-45 (in Chinese with English abstract) [本文引用:1]
47	王震宇, 张春荣, 李玲. 细胞分裂素结合蛋白的研究进展. 植物学通报, 2000, 17: 520-527 Wang Z Y, Zhang C R, Li L. Progress of studies on cytokinin-binding proteins. Chin Bull Bot, 2000, 17: 520-527 (in Chinese with English abstract) [本文引用:1]
48	董明辉, 刘晓斌, 陆春泉, 赵步洪, 杨建昌. 外源ABA和GA对水稻不同粒位籽粒主要米质性状的影响. 作物学报, 2009, 35: 899-906 Dong M H, Liu X B, Lu C Q, Zhao B H, Yang J C. Effects of exogenous ABA and GA on the main quality characteristics of grains at different positions of panicle in rice. Acta Agron Sin, 2009, 35: 899-906 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.667]
49	常二华, 王朋, 唐成, 刘立军, 王志琴, 杨建昌. 水稻根和籽粒细胞分裂素和脱落酸浓度与籽粒灌浆及蒸煮品质的关系. 作物学报, 2006, 32: 540-547 Chang E H, Wang P, Tang C, Liu L J, Wang Z Q, Yang J C. Concentrations of cytokinin and abscisic acid in roots and grains and Its relationship with grain filling and cooking quality of rice. Acta Agron Sin, 2006, 32: 540-547 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.667]
50	Travaglia C, Reinoso H, Cohen A, Luna C, Tommasino E, Castillo C, Bottini R. Exogenous ABA increases yield in field-grown wheat with moderate water restriction. J Plant Growth Regul, 2010, 29: 366-374 [本文引用:1] [JCR: 1.99]
51	崔志青, 尹燕枰, 田奇卓, 王广昌, 孟范玉, 王平, 王振林. 喷施ABA对两个穗型不同小麦穗颈节伤流、穗部性状及产量的影响. 生态学报, 2011, 31: 1085-1092 Cui Z Q, Yin Y P, Tian Q Z, Wang G C, Meng F Y, Wang P, Wang Z L. Effects of spraying ABA on bleeding intensity in neck-panicle node, spike traits and grain yields of two different panicle-type winter wheat. Acta Ecol Sin, 2011, 31: 1085-1092 (in Chinese with English abstract) [本文引用:1]
52	赵步洪, 张洪熙, 朱庆森, 杨建昌. 两系杂交稻籽粒充实不良的成因及其与激素含量的关系. 中国农业科学, 2006, 36: 477-486 Zhao B H, Zhang H X, Zhu Q S, Yang J C. Causes of poor grain plumpness of two-line hybrids and their relationships to contents of hormones in the rice grain. Sci Agric Sin, 2006, 36: 477-486 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.889]

1996

0.0

1.667

Acta Agron Sin. 1996, 22(6):736 - 740

Preliminary Studies on the Accumulation of Grain Starch and Protein during Grain Filling in Wheat

春小麦籽粒灌浆过程淀粉和蛋白质积累规律的初步研究

Liu Xiaobing;Li Wenxiong

刘晓冰;李文雄

Heilongjiang lnstitute of Agricultural Modernization,Academia sinica,Harbin 150040

... 小麦籽粒中蛋白质的含量和组分对小麦的产量、营养品质及加工品质均有重要影响^[1] ...

2006

0.0

... 植物激素是植物自身合成的生长调节物质, 对植物的生长发育发挥重要的调控作用^[2] ...

2004

0.0

... 烯效唑拌种提高了小麦叶片全氮含量^[3] ...

2000

0.0

0.849

... 细胞分裂素可诱导与蛋白质合成有关基因的表达, 调节蛋白质的合成^[4] ...

2007

0.0

0.849

... 小麦籽粒中玉米素核苷(ZR)含量与蛋白质含量呈显著正相关^[5] ...

2000

0.0

1.667

Acta Agron Sin. 2000, 26(6):659 - 665

Studies on the Relationship between Endogenous Hormone of Leaf and Accumulation of Nitrogen, Phosphorus and Potassium of Plant as Well as the Cause of Boll Weig ht Raised in High Yield Cotton

高产棉花叶片内源激素与氮磷钾吸收积累的关系及其对棉铃

CHEN De-Hua;HE Zhong-Pei;HE Zhong-Pei;ZHOU Gui-Sheng;WU Yun-Ka ng

陈德华;何钟佩;徐立华;周桂生;吴云康

The Key Opening Lab. of Crop Cultivation and Physiology of Jiangsu Province , Yangzhou University, Yangzhou 225009

The comprehensive effects of chemical fertilizer and DPC regulation on the endog enous hormone contents of cotton leaves as well as the physiological and biochem ical metabolism of high yield cotton plants were studied. The results are as fol lows: The improved contents of IAA, GA3 as well as ZR and

本文通过肥控结合对高产棉叶内源激素的影响及其棉株的生理生化代谢研究表明：肥料和缩节安(DPC)化控配合能促进结铃期棉叶中IAA、 GA3和ZR含量的提高，降低ABA的含量，从而诱导根系氨基酸和蛋白质的合成，促进棉叶RuBPase和蔗糖转化酶活性提高，有利于光合增强和光合产物输出，促进棉株各器官对氮、磷、钾的吸收，

... ZR在盛花期能促进棉花对氮素的吸收, 而降低结铃期后棉株ABA含量有利于氮素的吸收利用^[6] ...

2010

0.0

... GA₃可降低大豆根系硝态氮含量, 对硝酸还原酶有一定抑制作用^[7] ...

2010

0.0

... GA₃对棉铃Bt蛋白含量的调节效应与棉铃所处发育阶段有关^[8] ...

2010

0.0

1.889

Sci Agric Sin. 2010, 43(12):2595 - 2602

Effects of Spraying ABA on Glutenin Fraction Content and GMP Size Distribution in Wheat Grain

喷施ABA对小麦籽粒谷蛋白组分含量及GMP粒度分布的影响

CUI Zhi-qing, HE De-xian, CAI Tie, WANG Cheng-yu, WANG Guang-chang, MENG Fan-yu,HAN Zhan-jiang, LI Na-na, WANG Zhen-lin

崔志青,贺德先,蔡铁,王成雨,王广昌,孟范玉,韩占江,李娜娜,王振林

(山东农业大学农学院/作物生物学国家重点实验室)

【Objective】 In order to improve nutrient quality of wheat ( Triticum aestivum L.), the glutenin fraction content and glutenin macropolymer (GMP) changes of wheat grain were investigated after spraying ABA, as glutenin fraction content and glutenin macropolymer (GMP) play key roles in grain quality of wheat. 【Method】 The changes of glutenin fraction content and GMP size distribution were analyzed and the effects of ABA treatments on the glutenin fraction content and GMP size distribution in grains of two wheat cultivars, Shannong 8355 and Shannong 15, which had high grain yields in fields, were evaluated. 【Result】 The results showed that the protein content and glutenin content of wheat grain were improved after spraying ABA(12 mg•kg-1, 45 mL•m-2)at late booting stage and grain formation stage, the contents of HMW or LMW-insoluble glutenin in both of the two cultivars were also increased at the same time and reached a significant level when compared with the CK, but spraying ABA at the late booting stage, the grain soluble glutenin content was less affected. The GMP content and yield were increased and the particle size distribution of GMP were changed by spraying ABA, but the performance was different in different cultivars and treatments. Correlation analysis showed that HMW-insoluble glutenin content, LMW-insoluble glutenin content, GMP content and GMP/Pr. had a negative correlation with small particle size (d<15 um) distribution of GMP. 【Conclusion】 ABA could regulate the glutenin fraction content and glutenin macropolymer (GMP) size distribution and improve nutrient quality of wheat.

【目的】探讨喷施外源激素ABA引起小麦籽粒谷蛋白组分及谷蛋白大聚合体(glutenin macro-polymer, GMP)粒度分布的变化,为激素调控改善小麦籽粒品质提供理论依据。【方法】以冬小麦品种山农8355和山农15为试验材料,通过大田试验研究喷施外源ABA对小麦籽粒谷蛋白组分含量及GMP粒度分布的影响。【结果】在孕穗末期和籽粒形成期喷施ABA(浓度12 mg•kg-1,用量45 mL•m-2)均能提高籽粒粗蛋白及谷蛋白含量,同时,籽粒中高、低分子量不溶性谷蛋白含量均有所增加。特别是在孕穗期喷施ABA,山农8355和山农15两品种籽粒中高、低分子量不溶性谷蛋白含量与对照相比均达到显著水平,但喷施ABA对籽粒中可溶性谷蛋白含量影响相对较小。不论是孕穗期还是籽粒形成期喷施ABA均能增加GMP含量与产量,并改变籽粒中GMP的粒度分布,尤其是对其体积及表面积分布影响相对较大。相关分析表明,高、低分子量不溶性谷蛋白、GMP含量及GMP/Pr.比值均与GMP小颗粒(d<15 um)粒度分布呈现负相关,与大颗粒(d≥15 um)粒度分布的关系呈正相关。【结论】喷施外源ABA可以改变籽粒中谷蛋白组分及GMP粒度分布,从而影响小麦籽粒品质。

... 外源ABA可增加籽粒中蛋白质含量, 改变谷蛋白组分含量^[9] ...

... 此外, 也证实ABA处理显著提高汶农6号强、弱势籽粒的麦谷蛋白含量^[9] ...

2008

0.0

0.55

... 研究表明, 持绿型作物叶片叶绿素含量高, 植株衰老过程中叶片失绿缓慢^[10], 选育保绿品种是提高作物产量的有效途径^[11,12] ...

2000

0.0

0.965

J Maize Sci. 2000, 8(4):41 - 44

Advances in Physiological Basis of Maize that Stay-green

保绿型玉米的营养生理研究进展

HEPing;JINJi-yun

何萍;金继运

本文综述了近年来发展较快的保绿型玉米营养生理的研究进展,包括玉米保绿性的标准、玉米保绿性与植株含水量的关系、玉米保绿性与叶面积功能期的关系、保绿型玉米的营养生理基础及影响叶片保绿性的基因与环境因子等.

... 研究表明, 持绿型作物叶片叶绿素含量高, 植株衰老过程中叶片失绿缓慢^[10], 选育保绿品种是提高作物产量的有效途径^[11,12] ...

2000

5.242

0.0

... 研究表明, 持绿型作物叶片叶绿素含量高, 植株衰老过程中叶片失绿缓慢^[10], 选育保绿品种是提高作物产量的有效途径^[11,12] ...

2003

5.242

0.0

J. Exp. Bot.. 2003, 54(386):1415-20

Physiological characterization of 'stay green' mutants in durum wheat.

Spano G, Di Fonzo N, Perrotta C, Platani C, Ronga G, Lawlor DW, Napier JA, Shewry PR

Experimental Institute of Crop Research, S.S 16 km 675-71100 Foggia, Italy. g.spano@unifg.it

Four mutants with delayed leaf senescence were selected from seed of durum wheat mutagenized with ethylmethane sulphonate. Changes in net photosynthetic rate, efficiency of photosystem II and chlorophyll concentration during the maturation and senescence of the flag leaves of both mutant and parental plants were determined under glasshouse conditions. The four mutant lines maintained photosynthetic competence for longer than the parental line and are therefore functionally 'stay green'. The mutant lines also had higher seed weights and grain yields per plant than the parental line.

PMID: 12709488

... 持绿型小麦绿叶面积大, 叶片光合能力强, 有较长的灌浆持续期^[13] ...

2006

0.0

0.95

Chin Sci Bull. 2006, 51(18):2154 - 2160

(延缓小麦衰老的结构和生物化学机制),(科学通报),2006,51:2154-2460

.The structure and physiological and biochemical mechanism of delaying senescence of wheat

延缓小麦衰老的结构和生物化学机制

(罗培高),(任正隆),(吴先华),(张怀渝),(张怀琼),(冯娟).(延缓小麦衰老的结构和生物化学机制),(科学通报),2006,51:2154-2460

罗培高(1);任正隆(1,2);吴先华(1);张怀渝(1);张怀琼(1);冯娟(2)

川农17是一个含1RS.1BL易位染色体的保绿高产小麦新品种, 能有效地延缓叶片衰老和维持开花后较长时期的高光合性能. 生理、生物化学和透射电子显微镜结果表明, 川农17在灌浆期的旗叶净光合速率、叶绿素含量、丙二醛含量、超氧化物岐化酶活性、过氧化氢酶活性, 叶绿体及其超微结构变化的时间进程等方面, 与对照品种绵阳11存在显著差异, 且保持较长时间的强光合反应能力和光合速率. 叶绿体发育的超微结构变化与叶片生理衰老变化紧密相关. 川农17在生理、生物化学和细胞结构之间的协调发展是其获得高籽粒产量的重要保障.

... 不同持绿型小麦灌浆进程以及产量形成的生物学机制有显著差异^[14,15] ...

2010

0.0

1.237

J Nucl Agric Sci. 2010, 24(1):149 - 153

TRANSPORTATION AND DISTRIBUTION OF¹⁴C-RESERVES ON DIFFERENT GENOTYPE WHEAT VARIETIES AFTER ANTHESIS

(李涛),(龚月桦),(吕金印),(刘普灵).(不同基因型小麦¹⁴C-储备物在花后的转运与分配规律).(核农学报),2010,24:149-153

1.College of Life Sciences, Northwest Science and Technology University of Agriculture and Forestry Shaanxi, Yangling, 712100；2. Institute of Soil and Water Conservation, Northwest Science and Technology University of Agriculture and Forestry, Shaanxi, Yangling, 712100

Transportation and distribution of¹⁴C-reserves among different stay-green wheat varieties were studied by labelled flag leaf with¹⁴CO₂before anthesis. The results showed that about 80%-90% of¹⁴C-reserves, which assimilated before anthesis, had been kept in the stems and rachises at anthesis, and only 10%-20% remained in the leaves; after anthesis, these reserves were transported to the kernels. At maturity,¹⁴C-reserves in the leaves were almost entirely exported, but about 60%-80%¹⁴C-reserves remained in the stems and rachises, and about 20%-40% was distributed into the kernels. The¹⁴C-reserves in stay-green wheat was transported early from the leaves and stems, and the stay-green wheat YM66 has stored 40% of¹⁴C-reserves in the kernels at mature stages, which was much more than that in the varieties of XY22 and WM6.

利用¹⁴CO₂对不同持绿性状的小麦品种进行花前示踪标记，研究持绿小麦的¹⁴C-储备物在花后的转运与分配规律。结果表明，小麦花前标记的¹⁴C-储备物在开花时80%～90%已储存于茎杆和颖壳中，仅有约10%～20%残留在叶片中；开花后这些储备物都向籽粒转运，成熟时籽粒中¹⁴C分配率约为20%～40%，茎杆与颖壳中含有约60%～80%，极少量残留在叶片中。持绿型小麦YM66的¹⁴C-储备物从叶片和茎杆中输出的快，成熟时籽粒含有40%的¹⁴C-储备物，这样比率高于对照品种XY22 和早衰品种WM6的¹⁴C-储备物。

... 不同持绿型小麦灌浆进程以及产量形成的生物学机制有显著差异^[14,15] ...

... 这进一步证实持绿型小麦品种由于叶片功能期较长, 籽粒灌浆时间长, 为籽粒灌浆提供充足的营养物质, 产量一般高于非持绿品种^[15,20,21] ...

1988

0.0

1.667

Acta Agron Sin. 1988, 14(3):182 - 193

GROWTH ANALYSIS ON THE PROCESS OF GRAIN FILLING IN RICE

水稻籽粒灌浆的生长分析

Zhu Qingsen; Cao Xianzu; Luo Yiqi

朱庆森;曹显祖; 骆亦其

Jiangsu Agricultural College; Yangzhou; Jiangsu Province

Richards equation W=A/(1+Be~(-kt)) 1/N was used to describe theweight increasing of the strong and the weak grains of six rice varieties(including hybrids).According to N values in the equation of theweak grain,the grain-filling processes of the six rice varieties fall intotwo types,one was synchron

用 Richards 方程 W=A/(1+Be~(-kt))1/N 配合农垦57号、汕优63号等6个品种(含杂种)强、弱势粒的籽粒增重资料。依弱势粒配得方程的 N 值,将此6个水稻品种的籽粒灌浆状况分为两个类型,即:01的异步灌浆型。并据籽粒生长速率曲线的两个拐点,将籽粒灌浆过程划分为:前,中(盛)、后(至90%A 止)3个期。同时导出:起始生长势 Ro=K/N,生

... 2 籽粒增重及灌浆参数的计算按朱庆森等^[16]介绍的方法, 利用Richard方程模拟小麦籽粒增重动态 ...

1985

0.0

... 7, 即为小麦籽粒粗蛋白质含量^[17] ...

... 参照何照范^[17]的方法分离和测定蛋白质组分 ...

2009

0.0

1.883

Plant Nutr Fert Sci. 2009, 15(1):24 - 31 DOI:10.11674/zwyf.2009.0104

Effect of phosphorus on activities of enzymes related to nitrogen metabolism in flag leaves and protein content of wheat grains

施磷对小麦旗叶氮代谢关键酶活性和子粒蛋白质含量的影响

WANG Ping¹, YIN Yan-ping¹, FU Guo-zhan^{^1，2}, GUO Ying¹, CAI Rui-guo^{^1，3},
LIANG Tai-bo¹, GENG Qing-hui¹, WU Yun-hai¹, WANG Zhen-lin^1*

王平¹，尹燕枰¹，付国占^1，2，郭营¹，蔡瑞国^1，3，
梁太波¹，耿庆辉¹，邬云海¹，王振林^1*

1 National Key Laboratory of Crop Biology, Agronomy College of Shandong Agricultural University, Key Laboratory of Wheat Cultivation Physiology and Genetic Improvement, Ministry of Agriculture, Taian, Shandong, 271018；
2 Agronomy College, Henan University of Science and Technology, Luoyang 471003,China；
3 Agronomy Department of Hebei Normal University of Science and Technology, Changli, Heibei 066600, China

Effect of phosphorus on the activities of enzymes related to nitrogen metabolism in flag leaves, and protein content in grains of wheat cultivars with different end-use quality were studied under field conditions. The studied enzymes include nitrate reductase (NR), glutamine synthetase (GS), endop eptidase (EP) and carboxyp eptidase (CP). Field experiment was conducted at the Experiment Station of Shandong Agricultural University in Taian, Shandong from 2005 through 2006. Two cultivars, strong gluten cultivar JM20 and weak gluten cultivar SN1391, were selected and three phosphorus application levels, P₂O₅ 0, 135 and 225 kg/ha were set up. Results indicated that the effects of phosphorus application rates on enzyme activities and protein content varied with wheat cultivars. Both the enzyme activities and protein content in strong gluten wheat JM20 were promoted with increasing of phosphorus application rate, but declined in the weak gluten wheat SN1391. The un-unique response of different wheat cultivars on phosphorus should be fully considered during the process of developing P application strategy.

在田间试验条件下，以强筋品种济麦20和弱筋品种山农1391为材料，研究了不同施磷水平对2类型品种小麦旗叶氮代谢关键酶活性和子粒蛋白质积累的效应。结果表明，在P2O5 0～135 kg /hm²范围内，施磷明显提高济麦20灌浆前期旗叶硝酸还原酶（NR）和谷氨酰胺合成酶（GS）活性；但却降低山农1391旗叶NR、GS活性。强筋品种济麦20在灌浆中、后期旗叶内肽酶（EP）、羧肽酶（CP）活性和蛋白质含量因施磷而提高；而弱筋品种山农1391的EP、CP活性和子粒蛋白质含量则降低。施磷使济麦20子粒清蛋白、醇溶蛋白和谷蛋白的含量提高，球蛋白含量有所降低；但弱筋品种山农1391子粒球蛋白、醇溶蛋白和谷蛋白的含量因施磷而降低，清蛋白含量未见显著变化。说明小麦施磷应考虑不同品质类型品种氮代谢及子粒蛋白质含量对施磷的反应。

... 按王平等^[18]的方法测定谷氨酰胺合成酶(GS)活性 ...

2011

0.0

1.889

Sci Agric Sin. 2011, 44(13):2673 - 2682 DOI:10.3864/j.issn.0578-1752.2011.13.005

Effects of Spraying Exogenous ABA or GA on the Endogenous Hormones Concentration and Filling of Wheat Grains

外源ABA和GA对小麦籽粒内源激素含量及其灌浆进程的影响

YANG Wei-bing; WANG Zhen-lin; YIN Yan-ping; LI Wen-yang; LI Yong; CHEN Xiao-guang; WANG Ping; CHEN Er-ying; GUO Jun-xiang; CAI Tie; NI Ying-li

杨卫兵; 王振林; 尹燕枰; 李文阳; 李勇; 陈晓光; 王平; 陈二影; 郭俊祥; 蔡铁; 倪英丽

山东农业大学农学院/作物生物学国家重点实验室,

【Objective】The objective of the experiment is to explore the relationship between exogenous ABA and GA to the levels of endogenous hormones in grains and the grain filling, and provide a theoretical basis for increasing grain weight under the regulation of exogenous hormone.【Method】The field experiments were carried out at Taian Experimental Station of Shandong Agricultural University, two cultivars Shannong 1391 and Gaocheng 8901 were planted in the farm, exogenous ABA or GA to the spikes were sprayed at initial grain filling stage, spikes were sampled after anthesis till maturity, and grains were partitioned into two groups, superior grains (the 1st and 2nd grains from the basal part of middle-spikelets ) and inferior grains (the 3rd and 4th grains from the basal part of middle spikelets or the 3rd grains or the 3rd, 4th and 5th grains).Then, taking the grains for the analysis of endogenous hormone levels and grain filling characteristics.【Result】Spraying exogenous ABA or GA enhanced the grain weight. Spraying exogenous ABA enhanced the weight of superior grains. Spraying exogenous GA enhanced the weight of superior and inferior grain. At the same time, spraying exogenous GA enhanced the levels of endogenous GA in grains. Spraying exogenous ABA enhanced the levels of endogenous ABA in grains. Spraying exogenous ABA or GA enhanced the IAA and CTK content at middle and later periods of grains filling. Grain filling characteristics suggest that spraying exogenous ABA or GA extended the grain filling stage. Spraying exogenous ABA enhanced the initial grain filling rate of superior grains. Spraying exogenous GA enhanced the grain filling rate at middle and later periods of grains filling. 【Conclusion】Spraying exogenous ABA or GA regulated the levels of endogenous hormones in grains, aslo the initial grain filling stage and duration of grains filling stage, and then, grain weight was regulated. Spraying exogenous ABA enhanced the grain weight by enhancing the weight of superior grain. Spraying exogenous GA enhanced the grain weight by enhancing the weight of superior and inferior grain.

【目的】探讨喷施外源脱落酸（ABA）或赤霉素（GA）对小麦籽粒内源激素含量及其灌浆进程的影响，为激素调控提高籽粒的粒重提供理论依据。【方法】试验选用山农1391和藁城8901两个品种，籽粒灌浆初期喷施ABA或GA 两种外源激素于穗部，定期取籽粒样品，用高效液相色谱法测定籽粒内源激素含量，用三次多项方程式对籽粒灌浆进程进行模拟并分析相关参数。【结果】喷施外源ABA或GA均能显著增加两个品种的粒重。外源ABA或GA对粒重的调控存在粒位效应。喷施ABA显著增加强势粒粒重，喷施GA使强势粒和弱势粒粒重均显著增加。喷施外源ABA显著提高籽粒ABA含量，喷施外源GA显著提高籽粒GA含量，喷施外源ABA或外源GA均显著提高灌浆中后期籽粒IAA和CTK的含量，尤其是喷施外源GA效果更为明显。喷施外源ABA或外源GA均能延长籽粒的灌浆持续期，喷施ABA显著提高灌浆初期强势粒的灌浆速率，而喷施GA显著提高灌浆中后期强势粒和弱势粒的灌浆速率。【结论】喷施外源ABA或外源GA通过改变籽粒内源激素水平，调节灌浆起始时间和籽粒灌浆持续期，进而调控籽粒粒重的形成。喷施ABA是通过提高强势粒粒重来增加粒重的，喷施GA是通过提高强势粒和弱势粒的粒重来增加粒重的。

... 采用高效液相色谱法(HPLC)测定籽粒中4种内源激素的含量^[19] ...

... 外源ABA能提高内源ABA的含量^[19], 这一点也在本试验的两个品种中得到验证 ...

2008

0.0

... 这进一步证实持绿型小麦品种由于叶片功能期较长, 籽粒灌浆时间长, 为籽粒灌浆提供充足的营养物质, 产量一般高于非持绿品种^[15,20,21] ...

2009

0.716

0.0

... 这进一步证实持绿型小麦品种由于叶片功能期较长, 籽粒灌浆时间长, 为籽粒灌浆提供充足的营养物质, 产量一般高于非持绿品种^[15,20,21] ...

2006

0.0

0.692

... 植物激素在作物生长发育中起着重要的调控作用, 同化物质的转运及在库中的积累转化都有激素参与调控^[22,23] ...

2006

1.67

0.0

Plant Growth Regul. 2006, 49(1):1 - 8 DOI:10.1007/s10725-006-0017-3

The Relationship between Grain Filling and Hormone Content as Affected by Genotype and Source–sink Relation

Feng Wang(1) Fangmin Cheng(1) Guoping Zhang(1)

1.Department of Agronomy, Zhejiang University, Huajia Chi Campus, 310029 Hangzhou, China

The difference among florets within a panicle in grain filling and four hormone contents was compared by using two rice varieties differing in panicle type, i.e. compact-panicle variety Xiushui 994 and loose-panicle variety Xiushui 11. Meanwhile the effect of changing source (leaf cutting) and sink (floret removal) on grain filling and four hormone (indole-3-acetic acid, gibberellin, zeatin riboside and abscisic acid)level dynamics in developing grains was also examined. The results showed that Xiushui 994 had greater variation in grain filling rate (GFR), grain weight, filled grain percent (FGP) and four hormone contents during grain filling than Xiushui 11. Moreover, the former variety also showed the greater difference in the time of having the highest hormone contents between superior and inferior florets. The change in source and sink size had great effect on grain filling process and dynamics of four hormone contents, and the extent of the effect varied with variety and floret type. For Xiushui 994, floret-removal treatment had little effect on grain weight and FGP of the superior florets, but caused a marked decrease in GFR and four hormone contents, and a delay in the time when the highest GFR and hormone contents occurred. In the source-reducing treatment, GFR, grain weight and FGP for both superior and interior florets were all decreased. For Xiushui 11, both treatments had little effect on grain development in terms of grain filling properties and hormone content change in the two types of florets. The results indicated that the grain filling of compact-panicle variety is more source-limited than that of loose-panicle one, and relative level of hormone in both superior and inferior florets determined development of inferior florets.

... 植物激素在作物生长发育中起着重要的调控作用, 同化物质的转运及在库中的积累转化都有激素参与调控^[22,23] ...

2006

6.736

0.0

New Phytol.. 2006, 169(2):223-36

Grain filling of cereals under soil drying.

Yang J, Zhang J

Key Laboratory of Crop Genetics and Physiology of Jiangsu Province, Yangzhou University, Yangzhou, Jiangsu, China.

Monocarpic plants require the initiation of whole-plant senescence to remobilize and transfer assimilates pre-stored in vegetative tissues to grains. Delayed whole-plant senescence caused by either heavy use of nitrogen fertilizer or adoption of lodging-resistant cultivars/hybrids that remain green when the grains are due to ripen results in a low harvest index with much nonstructural carbohydrate (NSC) left in the straw. Usually, water stress during the grain-filling period induces early senescence, reduces photosynthesis, and shortens the grain-filling period; however, it increases the remobilization of NSC from the vegetative tissues to the grain. If mild soil drying is properly controlled during the later grain-filling period in rice (Oryza sativa) and wheat (Triticum aestivum), it can enhance whole-plant senescence, lead to faster and better remobilization of carbon from vegetative tissues to grains, and accelerate the grain-filling rate. In cases where plant senescence is unfavorably delayed, such as by heavy use of nitrogen and the introduction of hybrids with strong heterosis, the gain from the enhanced remobilization and accelerated grain-filling rate can outweigh the loss of reduced photosynthesis and the shortened grain-filling period, leading to an increased grain yield, better harvest index and higher water-use efficiency.

PMID: 16411926

... 外源ABA和细胞分裂素参与调控籽粒灌浆及籽粒中的物质代谢^[24,25], ABA参与光合产物的合成和积累^[26] ...

1999

0.0

1.667

Acta Agron Sin. 1999, 25(3):341 - 348

Regulation of ABA and GA to the Grain Filling of Rice

ABA与GA对水稻籽粒灌浆的调控

Yang Jianchang;Wang Zhiqin;Zhu Qingsen;Su Baolin

杨建昌;王志琴;朱庆森;苏宝林

Agricultural College of Yangzhou University, Yangzhou, 225009

ABA (Abscisic acid) content (ng g-1 FW) and the ratio of ABA to GA (Gibberellin) in strong potential grains or in the grains of the combination with good plumpness were higher than those in weak potential grains or in the grains of the combination with poor plumpness at early filling stage. Spraying

灌浆初期籽粒中ABA(脱落酸)含量(ng g-1FW)和ABA与GA(赤霉素)的比值，强势粒高于弱势粒，籽粒充实度好的组合高于籽粒充实度差的组合。开花后2天，喷施外源ABA，籽粒中ABA含量和ABA/GA增大，喷施外源GA后籽粒中GA含量增加，ABA/GA减小。低浓度(15mg/L)ABA处理后，灌浆初期籽粒中ADPG焦磷酸酶和淀粉合成酶活性及淀粉含量增加，

... 外源ABA和细胞分裂素参与调控籽粒灌浆及籽粒中的物质代谢^[24,25], ABA参与光合产物的合成和积累^[26] ...

... GA对籽粒灌浆起调控作用, 且与粒重的决定密切相关^[25] ...

... 抑制一些水解酶的活性, 并促进蛋白质的合成和积累^[25] ...

2007

0.0

... 外源ABA和细胞分裂素参与调控籽粒灌浆及籽粒中的物质代谢^[24,25], ABA参与光合产物的合成和积累^[26] ...

2004

1.67

0.0

Plant Growth Regul. 2004, 44(1):25 - 32 DOI:10.1007/s10725-004-1880-4

Effects of exogenous ABA and cytokinin on leaf photosynthesis and grain protein accumulation in wheat ears culturedin vitro

Xie

Z J

,Jiang

,Dai

T B

,Jing

,Cao

W X

.Effects of exogenous ABA and cytokinin on leaf photosynthesis and grain protein accumulation in wheat ears cultured in vitro.Plant Growth Regul,2004,44:25-32

1.Nanjing Agricultural University
2.Horticultural Research Institute, Shanghai Academy of Agricultural Sciences

A detached culture system and steady-state¹⁵N labeling technique were used to study the effects of exogenous ABA and ZR on photosynthetic characteristics, nitrogen remobilization and the activities of key enzymes for nitrogen metabolism in detached wheat parts during grain protein accumulation. The differences in net photosynthetic rate, chlorophyll content (SPAD value) and soluble protein content in the flag leaves of detached culture system between the treatments of ABA and ZR showed that ABA facilitates the post-anthesis senescence course compared to the ZR treatment. The differences in the changes of¹⁵N amount in different organs in the detached culture system between the ABA and ZR treatments showed that nitrogen remobilization from vegetative organs to the grain is accelerated by the ABA treatment but is delayed by ZR. The activities of GS and GPT in grains treated with ABA were significantly higher than those with the control treatment at 5 DAC, but reduced significantly compared with control at 11 DAC. The two enzyme activities in grains were reduced significantly by ZR at 5 DAC and increased significantly at 11 DAC, compared with those treated with ABA. The above changes of enzyme activity showed that the ABA treatment hastens amino acid conversion into grains and protein accumulation in grains, whereas the ZR treatment delays these processes. A significant reduction in grain weight with ABA treatment is associated with the reduction of net photosynthesis, chlorophyll content, and soluble protein content in flag leaves. Compared with the control and ZR treatments, a significant increase in grain protein content with the ABA treatment may result from the accelerating effects of ABA on N remobilization, amino acid conversion into grains and protein accumulation in grains.

... 研究表明, ABA促进氮素向籽粒的转运, 提高籽粒的蛋白质含量^[27] ...

1997

0.0

0.445

... ABA促进贮藏物质的积累, 特别是贮藏蛋白的合成^[28,29], 贮藏蛋白的基因受ABA信号的诱导而提高表达水平^[30] ...

2002

8.455

0.0

Curr Opin Plant Biol. 2002, 5(1):26 - 32 DOI:10.1016/S1369-5266(01)00225-4

ABA and sugar interactions regulating development: cross-talk or voices in a crowd?

Finkelstein

R R

,Gibson

S I

.ABA and sugar interactions regulating development: cross-talkor voices in a crowd.Curr Opin Plant Biol,2002,5:26-32

Abstract

Plant growth and development are controlled by the concerted action of many signaling pathways that integrate information from environmental signals with that from developmental and metabolic cues. Physiological studies have demonstrated that abscisic acid and sugars have both similar and antagonistic effects on diverse processes, including seed development, germination, and seedling growth. Recent genetic studies have identified several loci that are involved in both sugar and hormonal responses. It is rarely clear whether these apparent linkages reflect direct or indirect interactions between sugar and hormone signaling pathways, but the identification of gene products that are encoded at these loci is allowing these possibilities to be tested.

... ABA促进贮藏物质的积累, 特别是贮藏蛋白的合成^[28,29], 贮藏蛋白的基因受ABA信号的诱导而提高表达水平^[30] ...

2001

0.0

... ABA促进贮藏物质的积累, 特别是贮藏蛋白的合成^[28,29], 贮藏蛋白的基因受ABA信号的诱导而提高表达水平^[30] ...

1993

0.0

... 也有研究认为ABA对贮藏蛋白合成的促进作用发生在转录水平上^[31] ...

2013

1.632

0.0

... 谷氨酰胺合成酶是合成蛋白质的关键酶之一, 喷施ABA提高了GS活性(图6), 因而有利蛋白质的合成, 这与Yang等^[32]的报道一致, 即ABA通过提高GS活性促进蛋白质的合成 ...

2004

0.0

1.667

Acta Agron Sin. 2004, 30(10):1047 - 1052

Effects of Plant Growth Regulation Substances on Photosynthetic Characteristics and Assimilates Transportation in Winter Wheat under Post-anthesis Drought and Waterlogging

花后干旱和渍水条件下生长调节物质对冬小麦光合特性和物质运转的影响

XIE Zhu-Jie;JIANG Dong;CAO Wei-Xing;DAI Ting-Bo;JING Qi

谢祝捷;姜东;曹卫星;戴廷波;荆奇

Key Laboratory of Crop Growth Regulation, Ministry of Agriculture, Nanjing Agricultural University, Nanjing 210095, Jiangsu

Two winter wheat (Triticum aestivum L.) varieties differing in grain protein content, Wanmai38 and Yangmai9, were cultured under drought (soil relative water content, SRWC= 45%－50%) and waterlogging after anthesis in pot experiment to investigate the effects of two growth regulation substances i.e.

以温室盆栽的皖麦38（高蛋白）、扬麦9号（低蛋白）两个冬小麦(Triticum aestivum L.) 品种为材料，在花后土壤干旱（土壤相对含水量 = 45%～50%）和渍水状况下，分别施用6-BA（0.05 mmol/L）和NaHSO3（2 mmol/L），分析了两种生长调节物质对小麦光合特性、不同器官中物质运转和籽粒蛋白质及淀粉品质的影响。结果表明：两种生

... 6-BA能提高小麦花后同化氮素对籽粒总氮的贡献率^[33] ...

2009

2.699

0.0

J. Plant Physiol.. 2009, 166(16):1775-85 DOI:10.1016/j.jplph.2009.05.007

Cytokinin-induced changes of nitrogen remobilization and chloroplast ultrastructure in wheat (Triticum aestivum).

Criado MV, Caputo C, Roberts IN, Castro MA, Barneix AJ

IBYF-CONICET, Universidad de Buenos Aires, Av. San Martín 4453, Buenos Aires, Argentina. criado@agro.uba.ar

Nitrogen (N) remobilization in wheat (Triticum aestivum) plants is crucial because it determines the grain protein concentration and the baking quality of flour. In order to evaluate the influence of cytokinins on N remobilization during N starvation, we analyzed various N remobilization parameters in wheat plants that were watered with 6-benzylaminopurine (BAP) either with or without KNO(3). Besides, the effects of BAP on protein synthesis were evaluated, and the size and ultrastructure of chloroplasts of BAP-treated plants were studied. BAP supply inhibited N remobilization of plants independently of N supply as shown by the increase in protein, Rubisco, chlorophyll, sugar and starch concentrations in the older leaves, the decrease in amino acid and sugar export to the phloem, and the decrease in protein, Rubisco and chlorophyll concentrations in the younger leaves. Besides, BAP supply increased nitrate reductase activity and decreased nitrate concentration, thus suggesting an increased assimilatory capacity. The increase in protein concentration could be explained mainly by a significant decrease in protein degradation and, to a lesser extent, by an increase in protein synthesis. Finally, an increase both in the size of the chloroplast and in the plastoglobuli and starch contents in BAP-supplied plants was observed. We propose that cytokinins retain the sink activity of the older leaves by inhibiting amino acid and sugar export to the phloem and stimulating assimilate accumulation in the chloroplasts of the older leaves. Besides, BAP may increase protein concentration of the older leaves both by decreasing protein degradation and maintaining protein synthesis even under stress conditions.

PMID: 19540618

... Criado等^[34]发现, 6-BA能够抑制叶片蛋白质的降解及促进蛋白质的合成 ...

1999

2.32

0.0

... Ananieva等^[35]研究表明, 6-BA可促进茭瓜子叶中蛋白质和RNA的合成 ...

2010

0.0

1.667

Acta Agron Sin. 2010, 36(10):1769 - 1776 DOI:10.3724/SP.J.1006.2010.01769

Accumulation Dynamics of Glutenin Subunits and Glutenin Polymer in Wheat Cultivars with Different Qualities during Grain Development

不同品质类型小麦籽粒麦谷蛋白亚基及谷蛋白聚合体形成和累积动态

GAO Wen-Chuan,MA Meng,WANG Ai-Na,ZHAO Hui-Xian^*

高文川,马猛,王爱娜,赵惠贤

College of Life Science, Northwest A&F University, Yangling 712100, China

The quality of wheat (Triticum aestivum L.) flour for end use depends on the visco-elastic properties of dough, which are influenced by quantity and quality of the gluten-forming storage proteins of wheat endosperm. Gliadin and glutenin are major components of gluten protein. Dough strength is determined by the capacity of glutenin subunits to form large polymers with different sizes, and the quantity and composition of glutenin polymer influence wheat dough properties. To date, there are few reports on accumulation dynamics of glutenin subunits and glutenin polymer in wheat cultivars with different qualities during grain development. In this study, three strong-gluten and four weak-gluten wheat cultivars with different HMW-GS compositions were used to analyze accumulation dynamics of glutenin subunits and glutenin polymer during grain development. The results showed that HMW-GS and LMW-GS of strong-gluten cultivars initially expressed at 9–12 d after flowering, which were earlier than those of weak-gluten cultivars (12–15 d after flowering). Expression time of glutenin subunits in wheat cultivars with strong-gluten was a little earlier than that in wheat cultivars with weak-gluten. The quantity of HMW-GS in each cultivar rapidly increased during the early period of grain-filling, and kept stable from 27 d after flowering to maturity, however, the quantity of LMW-GS accumulated slowly during the grain-filling period, reaching stable level at 30 d after flowering. In wheat cultivars with strong-gluten, TGP (total glutenin polymer) % and GMP (glutenin macropolymer) % rapidly increased at 12–21 d after flowering and slowly increased during 21–30 d after flowering, and then maintained at the stable level from 30 d after flowering to maturity. However, in weak-gluten cultivars, TGP% and GMP% rapidly increased at 12–24 d after flowing and decreased continuously from 27 d after flowering to maturity. The differences of expression profile and accumulation of glutenin subunits and glutenin polymer between wheat cultivars with strong-gluten and weak-gluten may be the key factors affecting strong-gluten or weak-gluten formation of wheat cultivars.

选用高分子量麦谷蛋白亚基(HMW-GS)组成不同的3个强筋和4个弱筋小麦品种，研究了其籽粒发育过程中麦谷蛋白亚基、谷蛋白聚合体的形成和累积动态。结果表明，强筋小麦籽粒HMW-GS和B区低分子量麦谷蛋白亚基(LMW-GS)从花后9~12 d天开始表达；而弱筋小麦从花后12~15 d开始表达，即强筋小麦麦谷蛋白亚基开始形成时间早于弱筋小麦。各品种的HMW-GS一旦形成，其累积速度较快，花后27 d基本达到稳定值，之后维持稳定量；LMW-GS形成后，累积较慢，直到花后30 d左右达到稳定量。3个强筋小麦品种籽粒灌浆期谷蛋白总聚合体百分含量(TGP%)和谷蛋白大聚合体百分含量(GMP%)累积动态趋势基本一致，即在花后12~30 d一直持续增加，花后30 d至成熟达到最大值并保持稳定水平。4个弱筋小麦TGP%和GMP%累积动态均表现为在花后12~24 d (灌浆早中期)形成和持续累积，花后24 d至成熟逐渐降低。麦谷蛋白亚基表达模式以及谷蛋白聚合体累积动态的差异可能是导致小麦强筋或弱筋品质形成的关键。

... 蛋白质的组分分为清蛋白、球蛋白、醇溶蛋白和麦谷蛋白^[36] ...

2006

0.0

0.916

... 研究表明, 6-BA处理小麦穗可提高小麦谷蛋白/醇溶蛋白比值^[37] ...

2007

0.0

0.951

... 籽粒灌浆过程中各内源激素的含量在品种间存在基因型差异^[38] ...

2009

0.0

1.889

Sci Agric Sin. 2009, 42(12):4383 - 4389

Effects of Different Exogenous Hormones on Physiological Characteristics of Main Stem Leaves at Flower and Boll Stage in Cotton

不同外源激素对花铃期棉花主茎叶生理特性的影响

ZHENG Sha-sha, SUN Chuan-fan, SUN Hong-chun, LIU Lian-tao, ZHAO Jin-feng, LI Cun-dong

郑莎莎,孙传范,孙红春,刘连涛,赵金峰,李存东

（河北农业大学农学院/河北省作物生长调控重点实验室）

【Objective】 The effects of spraying exogenous hormones(gibberellin-3, GA3; brassinolide, BR and 6-benzylaminopurine, 6-BA) at early (July 6) and full (July 17) flowering time on physiological characteristics of main stem functional leaves at flower and boll stage. 【Method】 Three exogenous hormone treatments were emploid, with two concentration levels each, i.e. GA3 10 and 30 mg•L-1, BR 0.01 and 0.1 mg•L-1, and 6-BA 10 and 100 mg•L-1,with spraying water as the control(CK). The contents of chlorophyll, soluble protein, malondialdehyde, soluble sugar, endogenous hormone contents and the activities of superoxide dismutase (SOD) and peroxidase (POD) were measured. 【Result】 The results showed that the treatment of 100 mg•L-1 6-BA significantly increased the contents of chlorophyll, soluble protein, soluble sugar and the activities of SOD and POD at full flowering stage. The treatments of 6-BA 100 mg•L-1 and GA3 30 mg•L-1 significantly increased the contents of GA3 and ZR, and the ratio of ZR (zeatin riboside) / ABA (abscisic acid ),GA3 /ABA, and (ZR+GA3)/ABA at the same stage. The content of MDA greatly decreased. The treatments of BR (0.01 mg•L-1) and GA3 (10 mg•L-1) also had a positive effect, but the effect was weaker. The treatments of GA3 30 mg•L-1 and 6-BA 100 mg•L-1 significantly increased the content of soluble protein, the ratio of GA3/ABA, GA3+ZR/ABA, while the content of MDA significantly decreased at full bolling stage.【Conclusion】 Under this experimental condition, spraying 6-BA 100 mg•L-1 and GA3 30 mg•L-1 had significantly effects on the physiological functions of main stem functional leaves, it could be used for delaying the senescence of cotton.

【目的】研究初花期（7月6日）、盛花期（7月17日）喷施不同浓度GA3（赤霉素）、BR（油菜素内酯）和6-BA（6苄基腺嘌呤）对棉花盛花期（7月17日）和盛铃期（7月31日）主茎功能叶生理特性的影响。【方法】试验设GA3、BR和6-BA 3种激素处理,每种激素设2种浓度,即GA3(10和30 mg•L-1）、BR(0.01和0.1 mg•L-1）和6-BA(10和100 mg•L-1）,以喷清水为对照。测定主茎功能叶内的可溶性糖、丙二醛、叶绿素、可溶性蛋白质、内源激素含量,SOD（超氧化物歧化酶）和POD（过氧化物酶）活性。【结果】100 mg•L-1的6-BA能够显著提高盛花期主茎功能叶叶绿素、可溶性蛋白质含量和SOD、POD活性,使MDA（丙二醛）含量显著降低。100 mg•L-1的6-BA和30 mg•L-1的GA3处理,使盛花期内源ZR（玉米素核苷）、GA3含量和ZR/ABA（脱落酸）、GA3/ABA、（ZR+GA3）/ABA值均显著高于对照。0.01 mg•L-1的 BR和10 mg•L-1的GA3处理,盛花期主茎功能叶生理活性也有所提高。100 mg•L-1的6-BA和30 mg•L-1的GA3处理也显著提高了棉花盛铃期主茎功能叶可溶性蛋白质含量、GA3/ABA和GA3+ZR/ABA值,并使丙二醛含量显著降低。【结论】在本试验条件下,叶面喷施100 mg•L-1的6-BA、30 mg•L-1的GA3能够显著提高棉花花铃期主茎功能叶的生理活性,为生产中采用生长促进类外源激素防止棉花过早衰老提供了依据。

... 外源激素通过改变内源激素水平调节植物内在生理代谢的各个环节^[39] ...

2012

5.242

0.0

J. Exp. Bot.. 2012, 63(9):3499-509 DOI:10.1093/jxb/ers148

Plant hormone interactions: innovative targets for crop breeding and management.

Wilkinson S, Kudoyarova GR, Veselov DS, Arkhipova TN, Davies WJ

Lancaster Environment Centre, Lancaster University, Lancaster, UK. sally.wilkinson@lancaster.ac.uk

Here we highlight how both the root and shoot environment impact on whole plant hormone balance, particularly under stresses such as soil drying, and relate hormone ratios and relative abundances to processes influencing plant performance and yield under both mild and more severe stress. We discuss evidence (i) that abscisic acid (ABA) and ethylene act antagonistically on grain-filling rate amongst other yield-impacting processes; (ii) that ABA's effectiveness as an agent of stomatal closure can be modulated by coincident ethylene or cytokinin accumulation; and (iii) that enhanced cytokinin production can increase growth and yield by improving foliar stay-green indices under stress, and by improving processes that impact grain-filling and number, and that this can be the result of altered relative abundances of cytokinin and ABA (and other hormones). We describe evidence and novel processes whereby these phenomena are/could be amenable to manipulation through genetic and management routes, such that plant performance and yield can be improved. We explore the possibility that a range of ABA-ethylene and ABA-cytokinin relative abundances could represent targets for breeding/managing for yield resilience under a spectrum of stress levels between severe and mild, and could circumvent some of the pitfalls so far encountered in the massive research effort towards breeding for increases in the complex trait of yield.

PMID: 22641615

... 植物激素含量变化由基因表达调控^[40], 作物的持绿特性由细胞核及细胞质基因控制^[41] ...

2002

3.658

0.0

Theor Appl Genet. 2002, 104(4):526 - 532 DOI:10.1007/s001220100750

Isolation, characterization, and mapping of the stay green mutant in rice

K.-W. Cha(1) Y.-J. Lee(1) H.-J. Koh(1) B.-M. Lee(2) Y.-W. Nam(3) N.-C. Paek(1)

1.School of Plant Science, Seoul National University, Suwon 441-744, Korea e-mail: heejkoh@snu.ac.kr Tel.: +82-31-290-2307, Fax: +82-31-290-2560 KR
2.Department of Plant Resources, Dongguk University, Seoul 100-715, Korea KR
3.Department of Life Science, Sogang University, Seoul 121-742, Korea KR

Leaf color turns yellow during senescence due to the degradation of chlorophylls and photosynthetic proteins. A stay green mutant was isolated from the glutinousjaponica rice Hwacheong-wx through N-methyl-N-nitrosourea mutagenesis. Leaves of the mutant remained green, while turning yellow in those of the wild-type rice during senescence. The stay green phenotype was controlled by a single recessive nuclear gene, tentatively symbolized assgr(t). All the phenotypic characteristics of the mutant were the same as those of the wild-type lines except for the stay green trait. The leaf chlorophyll concentration of the mutant was similar to that of the wild-type before heading, but decreased steeply in the wild-type during grain filling, while very slowly in the mutant. However, no difference in photosynthetic activity was observed between the stay green mutant and the yellowing wild-type leaves, indicating that senescence is proceeding normally in the mutant leaves and that the mutation affects the rate of chlorophyll degradation during the leaf senescence. Using phenotypic and molecular markers, we mapped thesgr(t)locus to the long arm of chromosome 9 between RFLP markers RG662 and C985 at 1.8- and 2.1-cM intervals, respectively.

... 植物激素含量变化由基因表达调控^[40], 作物的持绿特性由细胞核及细胞质基因控制^[41] ...

2000

4.134

0.0

Plant Cell Physiol.. 2000, 41(3):251-7

Gibberellin biosynthesis: its regulation by endogenous and environmental signals.

Yamaguchi S, Kamiya Y

Department of Botany, Duke University, Durham, North Carolina 27708-1000, USA.

The hormone gibberellin (GA) plays an essential role in many aspects of plant growth and development, such as seed germination, stem elongation and flower development. In recent years, exciting progress has been made in understanding how the biosynthesis of this hormone is regulated by endogenous and environmental factors. This has resulted from isolation of genes encoding enzymes involved in GA biosynthesis and metabolism, which also enabled us to manipulate the pathway by modifying the expression of these genes in transgenic plants. In addition, new GA response mutants provided information about how signaling components are involved in feedback regulation of the GA biosynthetic pathway.

PMID: 10805587

... 研究表明, GA的生物合成受GA含量的反馈调节, 低含量的GA会激活GA的生物合成^[42] ...

1999

11.808

0.0

Trends Plant Sci.. 1999, 4(12):472-478

Formation and breakdown of ABA.

Cutler AJ, Krochko JE

Plant Biotechnology Institute, National Research Council of Canada, Saskatoon, Canada S7N 0W9.

The phytohormone, abscisic acid (ABA) is found in all photosynthetic organisms. The amount of ABA present is determined by the dynamic balance between biosynthesis and degradation: these two processes are influenced by development, environmental factors such as light and water stress, and other growth regulators. ABA is synthesized from a C40 carotenoid precursor and the first enzyme committed specifically to ABA synthesis is a plastid- localized 9-cis-epoxycarotenoid dioxygenase, which cleaves an epoxycarotenoid precursor to form xanthoxin. Subsequently, xanthoxin is converted to ABA by two cytosolic enzymes via abscisic aldehyde, but there appears to be at least one minor alternative pathway. The major catabolic route leads to 8'-hydroxy ABA and phaseic acid formation, catalyzed by the cytochrome P450 enzyme ABA 8'-hydroxylase. In addition, there are alternate catabolic pathways via conjugation, 4'-reduction and 7'-hydroxylation. As a consequence of recent developments, the mechanism by which the concentration of hormonally active ABA is controlled at the cellular, tissue and whole plant level can now be analyzed in detail.

PMID: 10562731

... ABA的生物合成类萜途径是由甲瓦龙酸(MVA)经中间产物异戊烯基焦磷酸(IPP)等一系列步骤^[43,44], 而IPP也是玉米素生物合成的起始物^[45], 因而本试验中喷施ABA提高ZR含量可能是通过促使IPP向玉米素合成实现的, 而外源ABA提高GA₃含量可能是通过降低前期内源GA₃含量, 由GA₃的反馈调节提高籽粒灌浆后期的GA₃含量实现的 ...

2002

11.808

0.0

Trends Plant Sci.. 2002, 7(1):41-8

Complex regulation of ABA biosynthesis in plants.

Seo M, Koshiba T

Dept of Biological Sciences, Tokyo Metropolitan University, Hachioji-shi, 192-0397, Tokyo, Japan. sseo@hoshoku-mac2.biol.metro-u.ac.jp

Abscisic acid (ABA) is a plant hormone that plays important roles during many phases of the plant life cycle, including seed development and dormancy, and in plant responses to various environmental stresses. Because many of these physiological processes are correlated with endogenous ABA levels, the regulation of ABA biosynthesis is a key element facilitating the elucidation of these physiological characteristics. Recent studies on the identification of genes encoding enzymes involved in ABA biosynthesis have revealed details of the main ABA biosynthetic pathway. At the same time, the presence of gene families and their respective organ-specific expression are indicative of the complex mechanisms governing the regulation of ABA biosynthesis in response to plant organ and/or environmental conditions. There have been recent advances in the study of ABA biosynthesis and new insights into the regulation of ABA biosynthesis in relation to physiological phenomena.

PMID: 11804826

2006

0.0

2005

0.0

... 内源细胞分裂素可以提高籽粒灌浆强度, 增大千粒重^[46] ...

2000

0.0

... 细胞分裂素与其特定受体结合后诱导基因表达, 促进RNA和蛋白质的合成^[47] ...

2009

0.0

1.667

Acta Agron Sin. 2009, 35(5):899 - 906 DOI:10.3724/SP.J.1006.2009.00899

Effects of Exogenous ABA and GA on the Main Quality Characteristics of Grains at Different Positions of Panicle in Rice

外源ABA和GA对水稻不同粒位籽粒主要米质性状的影响

DONG Ming-Hui¹,LIU Xiao-Bin²,LU Chun-Quan¹,ZHAO Bu-Hong²,YANG Jian-Chang^1*

董明辉¹，刘晓斌²，陆春泉¹，赵步洪²，杨建昌^1,*

1Fostering Base for the National Key Laboratory of Crop Genetics and Physiology of Jiangsu Province,Yangzhou University,Yangzhou 225009,China;2Lixiahe Region Agricultural Research Institute of Jiangsu,Yangzhou 225007,China

Plant hormones play important roles in plant growth and development and yield formation. ABA and GA are two important kinds of hormones controlling rice grain development and filling, but the mechanism of regulating grain quality by them is not understood completely. In order to study and document the influence of ABA and GA on grain quality, the solutions with low concentrations of exogenous ABA (75.7 μmol L^-1) and GA(57.7 μmol L^-1) were sprayed at earlier filling stage. The results indicated that the effects of exogenous hormones at early filling stage on rice quality were great, and varied with exogenous hormone varieties and different grain positions. The exogenous ABA increased the 1000-grain weight (KGW) and head milled rice rate (HMRR), and reduced chalkiness degree (CD); the effect of ABA on grain quality varied with the positions of branch and grain in a rice panicle, which was greater on the later-flowered spikelets than on the earlier-flowered spikelets in the same branch, and on second branch than on primary branch. With spraying exogenous GA, KGW, and HMRR, gel consistence and crude protein content (CPC) were decreased significantly, while CD and amylase content were increased. The effect of spraying GA on KGW and HMRR of the earlier-flowered spikelets was greater than that of the later-flowered spikelets, but it was reversed on CD, AC, and CPC compared with spraying ABA.

以中熟籼稻扬稻6号和中熟粳稻扬粳9538为材料，灌浆初期通过喷施低浓度的ABA (75.7 μmol L^-1)和GA (57.7 μmol L^-1)处理，研究其对水稻穗部不同粒位籽粒品质的影响。结果表明，灌浆初期喷施低浓度的外源ABA和GA对米质影响较大，影响程度因激素种类和籽粒着生部位不同而异。ABA增加千粒重和整精米率，降低垩白度；其影响对同枝梗上开花较迟的籽粒大于对其他粒位籽粒，对同部位二次枝梗籽粒大于对一次枝梗籽粒。外源GA处理则显著或极显著降低千粒重、整精米率、胶稠度、粗蛋白含量，增加垩白度、直链淀粉含量；同一枝梗不同粒位籽粒间，对较早开花籽粒千粒重和整精米率的影响大于对较迟开花籽粒，对垩白度、直链淀粉含量和蛋白质的影响则与ABA相反。

... -淀粉酶等水解酶而不利于淀粉和蛋白质的合成和积累^[48] ...

2006

0.0

1.667

Acta Agron Sin. 2006, 32(4):540 - 547

Concentrations of Cytokinin and Abscisic Acid in the Roots and Grains and Relationship with Grain Filling and Cooking Quality of Rice

水稻根和籽粒细胞分裂素和脱落酸浓度与籽粒灌浆及蒸煮品质的关系

CHANG Er-Hua; ZHANG Wen-Jie; TANG Cheng; LIU Li-Jun; WANG Zhi-Qin and YANG Jian-Chang

常二华;张文杰;唐成 ;刘立军;王志琴;杨建昌

Key Laboratory of Crop Genetics and Physiology of Jiangsu Province, Yangzhou University, Yangzhou 225009, Jiangsu，China

Cytokinin and abscisic acid (ABA) are two kinds of important plant hormones. Their roles in the grain filling and quality of rice, however, are not fully understood. The objective of this study was to determine changes in concentrations of zeatin (Z) + zeatin riboside (ZR) and ABA in both the roots and grains during grain filling, grain filling characteristics and the indexes for cooking quality using ten rice cultivars (including hybrids and lines) grown in nutrient solution. The results showed that the change in concentrations of Z+ZR and ABA in roots and grains exhibited a single peak curve during the grain filling period. The Z + ZR concentration in roots was very high at early grain filling stage and reached a peak at 3–9 d after anthesis (DAA), and that in grains at 9–12 DAA. ABA concentrations in roots and grains were rather low at early grain filling stage, then increased with grain filling, and reached a peak at 15–18 DAA in roots and at 15–21 DAA in grains. Concentrations of Z + ZR in roots and grains at early grain filling stage (0–12 DAA) and ABA at the mid stage (13–26 DAA) were significantly or very significantly correlated with the initial grain growth potential, mean grain filing rate, maximum grain filling rate and brown rice weight (r=0.726* ~ 0.984**), whereas significantly and negatively correlated with active grain filling period (r=－0.749* ~ －0.834**). Z+ZR concentrations at the mid or late grain filling stage (27–40 DAA) significantly or very significantly correlated with active grain filling period (r=0.689* ~ 0.932**) but those at the late stage negatively correlated with grain filling rate (r=－0.826** ~ －0.927**). The gel consistency and alkali spreading value of rice were significantly or very significantly correlated with Z+ZR concentrations in roots and grains at mid and late grain filling stages (r = 0.722* ~ 0.896**), whereas significantly and negatively correlated with ABA concentrations at mid grain filling stage (r=－0.883**~－0.913**). The correlations between amylose content and the concentrations of Z+ZR and ABA were reversed to the correlations of gel consistency with the two hormones. When roots were treated with 10^-7 mol/L ZR or with 10^-7 mol/L ABA at 0-6, 14–20 and 24–30 DAA, respectively, the effects of the exogenous hormones on grain filling and the cooking quality were consistent with the relationships of the two endogenous hormones with the grain filling parameters and indexes for cooking quality. The results suggest that cytokinin and ABA play important roles in the regulation of grain filling and quality of rice, and the direction of effects may dependent on grain filling stages.

以10个不同基因型水稻品种和杂交稻组合为材料，分析了结实期水稻根系和籽粒细胞分裂素和脱落酸（ABA）浓度的变化及其与籽粒灌浆速率和稻米蒸煮品质的关系。结果表明，灌浆早期(花后0～12 d)根和籽粒玉米素＋玉米素核苷（Z＋ZR）浓度以及灌浆中期（花后13～26 d）根和籽粒ABA浓度与籽粒起始生长势、平均灌浆速率、最大灌浆速率、糙米重呈显著或极显著正相关（r＝0.726* ~ 0.984**），与活跃灌浆期呈显著或极显著负相关(r＝－0.749* ~ －0.834**)。灌浆中、后期（花后27~40 d）根和籽粒Z＋ZR浓度与活跃灌浆期呈显著或极显著正相关(r＝0.689* ~ 0.932**)，但灌浆后期Z＋ZR浓度与灌浆速率呈极显著负相关(r＝－0.826** ~ －0.927**)。灌浆中期和后期根和籽粒Z＋ZR浓度与胶稠度及碱化值呈显著或极显著正相关（r＝0.722* ~ 0.896**），与直链淀粉含量呈显著或极显著负相关（r＝－0.633* ~－0.778**）。灌浆中期根和籽粒ABA浓度与胶稠度及碱化值呈极显著负相关（r＝－0.883** ~ －0.913**），与直链淀粉含量呈显著或极显著正相关（r＝0.803** ~ 0.871**）。不同灌浆期ZR或ABA处理对籽粒灌浆、稻米胶稠度和直链淀粉含量的影响，与内源激素同灌浆特征参数和稻米蒸煮品质指标的关系基本吻合。表明根和籽粒细胞分裂素和ABA对籽粒灌浆和稻米蒸煮品质起调控作用，其调控的正、负效应取决于灌浆的时期。

... ABA表现多种生理功能, 如促进水稻籽粒灌浆^[49] ...

2010

1.99

0.0

J Plant Growth Regul. 2010, 29(3):366 - 374 DOI:10.1007/s00344-010-9147-y

Exogenous ABA Increases Yield in Field-Grown Wheat with Moderate Water Restriction

Claudia Travaglia(1) Herminda Reinoso(1) Ana Cohen(2) Celina Luna(3) Ezequiel Tommasino(3) Carlos Castillo(4) Rubén Bottini(2)

1.Departamento de Ciencias Naturales, Facultad de Ciencias Exactas, Físico, Químicas y Naturales, Universidad Nacional de Río Cuarto, 5800 Río Cuarto, Argentina
2.Cátedra de Química Orgánica y Biológica, Facultad de Ciencias Agrarias, Universidad Nacional de Cuyo, 5505 Chacras de Coria, Mendoza Argentina
3.Instituto de Fitopatología y Fisiología Vegetal (IFFIVE) INTA, 5119 Córdoba, Argentina
4.área de Producción de Cereales, Facultad de Agronomía y Veterinaria, Universidad Nacional de Río Cuarto, 5800 Río Cuarto, Argentina

Water stress is one of the most important environmental factors that regulate a plant’s growth and development. In agronomic practice the effects of water stress are translated into low yield and/or reduced quality. Abscisic acid (ABA) sprays (1 mM) were applied to wheat plants at different phenological stages and the effects on several physiological variables and on yield were evaluated under field conditions at different water regimes. Studies were conducted in the field across three consecutive winter–spring seasons. ABA treatments were applied at the beginning of shoot enlargement and repeated at anthesis. Exogenous ABA increased shoot dry weight and maintained a high concentration of photosynthetic pigments for a longer period of time during grain growth and maturation. Although ABA applications increased stomatal closure immediately after its application, the longer-term effect was to allow for a greater ostiolar opening of the stomatal pore which resulted in increased conductance of gases and water vapor. ABA also improved the transport of photoassimilates from the leaves and stem to the developing grains, that is, it effectively increased the sink strength of the grains. This correlated with a yield increase without significantly changing the protein quality in the grains. Thus, elevated ABA levels from exogenous application or genetic selection could help improve agricultural production of grains in arid areas where irrigation is not possible.

... 延长小麦叶片保绿时间, 延缓叶绿素降解^[50] ...

2011

0.0

... 提高小麦穗颈节伤流强度, 促进营养物质的运转^[51] ...

2006

0.0

1.889

... IAA对胚乳细胞的发育以及籽粒中物质积累有调控作用^[52] ...