张雪, 杨洪坤, 郑亭, 肖云, 莫飘, 樊高琼^,四川农业大学农学院/农业部西南作物生理生态与耕作重点实验室,成都 611130

Effects of Exogenous ABA on Pre-Harvest Sprouting Resistance and Quality of White and Red Wheat Cultivars

ZHANG Xue, YANG HongKun, ZHENG Ting, XIAO Yun, MO Piao, FAN GaoQiong^,College of Agronomy, Sichuan Agricultural University/Key Laboratory of Crop Eco-Physiology and Farming System in Southwest China, Ministry of Agriculture, Chengdu 611130

通讯作者: 樊高琼,E-mail： fangao20056@126.com

责任编辑: 杨鑫浩
收稿日期:2020-02-24接受日期:2020-06-23网络出版日期:2020-12-01

基金资助:

国家重点研发计划项目.2016YFD0300406 四川省农作物育种攻关项目.2016NYZ005 国家公益性行业(农业)科研专项.20150312705

Received:2020-02-24Accepted:2020-06-23Online:2020-12-01
作者简介 About authors
张雪,E-mail： sdz626zx@163.com。








摘要
【目的】西南麦区收获季多阴雨导致籽粒易穗发芽是造成该区域小麦品质变劣和商品性差的重要影响因素。研究脱落酸（ABA）喷施时期和喷施浓度对小麦穗发芽抑制效果及其对小麦籽粒品质的影响,提出适宜该区域的ABA喷施组合技术模式,为生产上利用植物生长调节剂调控穗发芽和改善小麦品质提供理论依据与技术支撑。【方法】以易穗发芽白皮小麦品种中科麦138和抗穗发芽红皮小麦品种绵麦367为试验材料,在灌浆初期（15 DAA）、灌浆后期（30 DAA）以及生理成熟期（35DAA）,喷施不同浓度ABA（0、50、100 mg·L ^-1）,研究其对两粒色小麦品种穗发芽表型,灌浆期间α-淀粉酶活性、籽粒可溶性糖和淀粉含量动态,收获后籽粒蛋白质、湿面筋含量、沉淀值,淀粉组分及RVA特征值的影响。【结果】（1）花后喷施不同浓度ABA对小麦穗发芽均有抑制作用,2个年度均以花后30 d喷施抑制发芽效果最好;收获期雨水较少年份（2018年）施用50 mg·L ^-1喷施浓度即可,中科麦138生理成熟期及蜡熟期的粒发芽率较对照下降13.8和3.8个百分点,绵麦367则较对照下降23.5和9.7个百分点;收获期雨水较多年份（2019年）则以100 mg·L ^-1喷施浓度抑制作用更优,中科麦138生理成熟期及蜡熟期的粒发芽率较对照下降22.5和19.6个百分点,绵麦367则较对照下降10.0和12.0个百分点;同时,不同时期不同浓度ABA处理后对种子发芽的抑制作用均在收获后60 d得以解除,不影响后续正常发芽。（2）外源喷施ABA后可降低α-淀粉酶活性,对穗发芽敏感期（花后35—45 d）α-淀粉酶活性抑制作用显著,以花后30 d喷施抑制效应最好,该时期喷施100 mg·L ^-1ABA,花后45 d籽粒α-淀粉酶活性较对照下降30.1%,可溶性糖含量较对照下降41.9%,而淀粉含量较对照提高10.2个百分点,淀粉水解受到抑制。（3）喷施ABA后可提高蛋白质质量,100 mg·L ^-1喷施浓度处理的沉淀值较CK提高4.3%—8.8%;外源喷施ABA对籽粒淀粉组分及面粉糊化特性影响更大,处理后支链淀粉含量增加进而总淀粉含量增加;100 mg·L ^-1喷施浓度处理的支链淀粉和总淀粉含量分别较CK增加8.1和7.6个百分点,直/支比下降18.2%,面粉糊化特性进一步改善,降落值、峰值粘度和崩解值提升,随浓度增大呈增加趋势,100 mg·L ^-1喷施浓度处理的降落值、峰值粘度和崩解值较CK提高幅度分别为20.9%—24.2%、26.5%—51.4%和12.4%—43.4%。【结论】西南麦区于花后30 d喷施50—100 mg·L ^-1ABA,可有效抑制穗发芽敏感期α-淀粉酶活性,抑制淀粉水解,降低穗发芽率和粒发芽率,提高蛋白质质量,并可增加支链淀粉含量和总淀粉含量,降低直/支比,改善面粉的糊化特性,可作为西南麦区生育后期增强小麦穗发芽抗性及减损提质的重要栽培管理措施,建议加大其示范与推广力度。
关键词： ABA;小麦;穗发芽;籽粒品质

Abstract
【Objective】Pre-harvest sprouting (PHS) resulted from higher rainfall during the harvest period is one of the crucial reasons degrade food uses quality of wheat flour in Southwest wheat production region of China. This study was carried out to investigate the inhibition effect of exogenous abscisic acid (ABA) to pre-harvest sprouting, and the changes in grain quality with respecting to PHS sensitivity was also measured. 【Method】Two cultivators with pre-harvest sprouting resistance (sensitive: white-seeded wheat Zhongkemai 138; insensitive: red-seeded wheat Mianmai 367) were used as experimental materials. The anti-photodegraded ABA (0, 50, and 100 mg·L ^-1) were sprayed at 15 (initial filling stage), 30 (late filling stage) and 35 days (physiological maturity stage) after anthesis (DAA) to investigate the changes in germination traits, α-amylase activity and grain quality. 【Result】 (1) The application of 50-100 mg·L ^-1 ABA at 30 DAA could inhibit PHS, with the optimum spraying time achieved at 30 DAA. In normal years (2018), the wheat spraying with 50 mg·L ^-1ABA was better than other treatments as compared to the control, the germination rate of Zhongkemai 138 was reduced by 13.8 and 3.8 percentage points at physiological maturity and dough stage, respectively, and the PHS sensitivity cultivator (Mianmai 367)was decreased by 23.5 and 9.7 percentage points, respectively, as compared with control. In 2019 (rainy season), spraying with 100 mg·L ^-1 ABA performed better than other treatments with the germination rate of Zhongkemai 138 was reduced by 22.5 and 19.6 percentage points during the physiological maturity and dough stage, respectively, as compared with control, and PHS sensitivity cultivator (Mianmai 367) was reduced by 10.0 and 12.0 percentage points than that of control, respectively. Meanwhile, the inhibitory effects of ABA were all released at 60 days after harvest, and did not affect subsequent seed germination. (2) Exogenous application of ABA could reduce α-amylase activity and inhibited α-amylase activity, which further delayed the hydrolysis of starch in 35-45 DAA. Compared with the control, the α-amylase activity and soluble sugar content were decreased by 30.1%, and 41.9%, respectively, and the starch content was 10.2 percentage points higher when the 100 mg·L ^-1 ABA was spraying at 30 DAA. (3) The application of 100 mg·L ^-1 ABA improved the precipitation value by 4.3%-8.8%, and exogenous application of ABA showed a greater impact on the starch content with the amylopectin content and total starch content increased by 8.1 and 7.6 percentage points, and the amylose/amylopectin ratio decreased by 18.2%. Further, the pasting properties of flour were also improved with the falling value, peak viscosity and disintegration value was 20.9%-24.2%, 26.5%-51.4% and 12.4%-43.4% higher than that of CK, respectively. 【Conclusion】The application of 50-100 mg·L ^-1 ABA at 30 DAA could effectively reduce pre-harvest sprouting by inhibiting α-amylase induced starch hydrolysis without decline the protein quality, and thereby improved the pasting properties of wheat flour by enhancing amylopectin and total starch contents, reducing the ratio of amylase and amylopectin. Therefore, the application of 50-100mg·L ^-1 ABA at 30 DAA was highly recommended for farmers to enhance the pre-harvest sprouting resistance and reducing the losses in food uses quality in Southwest wheat production region of China.
Keywords：abscisic acid;wheat;pre-harvest sprouting;grain quality


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本文引用格式
张雪, 杨洪坤, 郑亭, 肖云, 莫飘, 樊高琼. 外源ABA对两粒色小麦品种穗发芽及品质的影响[J]. 中国农业科学, 2020, 53(23): 4750-4763 doi:10.3864/j.issn.0578-1752.2020.23.003
ZHANG Xue, YANG HongKun, ZHENG Ting, XIAO Yun, MO Piao, FAN GaoQiong. Effects of Exogenous ABA on Pre-Harvest Sprouting Resistance and Quality of White and Red Wheat Cultivars[J]. Scientia Agricultura Sinica, 2020, 53(23): 4750-4763 doi:10.3864/j.issn.0578-1752.2020.23.003


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0 引言

【研究意义】小麦穗发芽（pre-harvest sprouting,PHS）是世界性的自然灾害^[1,2],穗发芽消耗籽粒内部贮藏物质,破坏籽粒形态,造成产量下降,品质劣化,严重影响小麦经济价值^[3,4]。西南麦区作为我国第三大小麦优势产区,在小麦收获期常遭遇连阴雨天气,小麦穗发芽现象尤为普遍和严重^[5,6],穗发芽是导致该区域小麦品质次、商品性差的重要原因之一,如何有效预防或降低穗发芽已成为制约该区小麦产业发展的重大问题。【前人研究进展】穗发芽是困扰全世界小麦育种家100多年的大难题,由于抗源缺乏,抗穗发芽育种进展缓慢^[7]。前人多从种子休眠的激素响应研究穗发芽^[8,9],调控措施上,往往通过施用休眠促进剂（脱落酸、多效唑、烯效唑）改变ABA/GA比例,降低α-淀粉酶活性,延长种子休眠而抑制穗发芽发生^[10,11,12]。有关ABA施用时期和浓度,不同作物上报道不同。如雍太文等^[13]发现开花期喷施90 mg·L^-1ABA或者灌浆中期喷施50 mg·L^-1 ABA均能有效抑制水稻穗发芽。杨光宇等^[14]在花后36 d喷施80 mg·L^-1穗发芽抑制剂也能有效抑制小麦穗发芽。另有研究表明,ABA在稻麦籽粒灌浆及品质形成中亦具有重要调控作用^{[15,16,17,18]}。YANG等^[19]研究认为,在灌浆初期水稻籽粒中ABA含量与细胞分裂速率呈负相关,而在籽粒生长的线性阶段（11—15 DAA）施用ABA可显著提高籽粒灌浆速率和粒重。戴忠民等^[17]研究结果表明,花后4 d喷施ABA可显著降低胚乳细胞增殖速率,减少强势籽粒的胚乳细胞数,且能降低直链淀粉含量及淀粉直/支比。崔志青等^[20]研究结果认为,小麦孕穗末期和籽粒形成期喷施ABA,籽粒粗蛋白及谷蛋白含量均有所提高。【本研究切入点】前人有关ABA对萌发和休眠调控的基础研究较多,有关ABA不同时期喷施在灌浆中的作用及对籽粒品质的影响也有较多报道,但有关ABA作为穗发芽抑制剂报道较少,研究较浅,喷施时期和浓度各异,且生产应用少见;同时,ABA抑制穗发芽效应与品质效应分开而论,与当今的生产问题和市场需求脱节。【拟解决的关键问题】本研究通过设置外源ABA喷施时期与喷施浓度,对两粒色小麦品种穗发芽及品质的调控主效应及其互作效应的影响进行研究,旨在明确ABA抑制小麦穗发芽的最佳施用时期和浓度,为生产中小麦后期抗灾、减损、提质化控技术的制定提供理论依据和技术支撑。

1 材料与方法

1.1 试验设计

试验于2017—2019年在四川省大邑县安仁镇现代农业（粮食产业）园区进行,试验田位于成都平原西部,属亚热带湿润季风气候。供试土壤为水稻土,土壤0—20 cm土层有机质含量14.5 g·kg^-1,碱解氮184.1 mg·kg^-1,速效钾142.2 mg·kg^-1,前茬为水稻。试验采用三因素裂-裂区设计,品种为主区,喷药时期为裂区,ABA浓度为裂裂区。供试品种为白皮小麦品种中科麦138（ZKM138,易穗发芽）和红皮小麦品种绵麦367（MM367,抗穗发芽）,分别由中国科学院成都生物研究所和绵阳农业科学研究院提供;ABA悬浮剂（有效成分含量5%,抗光解）为中国农业大学作物化控创新研究团队提供。2017—2018年于灌浆后期（B₁,花后30 d）和生理成熟期（B₂,花后35 d）进行穗部喷施。第1年的结果表明,B₂时期喷施对穗发芽抑制效应显著低于B₁时期喷施,喷施时期是否需要提前到灌浆初期?基于此,2018—2019年度施药时期保留B₁（花后30 d）,删去B₂,增加B₃（灌浆初期即花后15 d）和B₄（花后15 d与花后30 d叠加,即花后（15+30）d））。2个年度ABA喷施浓度均为50 mg·L^-1（C₁）和100 mg·L^-1（C₂）,以喷等量清水为对照（CK）,每个喷药时期均连续喷施2 d,1次/d,每次喷药量为600 kg·hm^-2,喷施时间为17：00—18：00。

2个年度均于10月30日采用免耕撬窝点播,行距20 cm,穴距10 cm,小区面积为8 m²（2 m×4 m）,3次重复。基本苗控制在2.4×10⁶ hm^-2;全生育期施150 kg N·hm^-2、75 kg P₂O₅·hm^-2和75 kg K₂O·hm^-2,其中60%的氮肥和全部磷钾肥用作底肥,40%氮肥于拔节期追施,其他栽培措施按大田生产进行。

2年小麦灌浆期间降雨情况如图1所示,2017—2018年度花后35—45 d基本无降雨,小麦正常收获;而2018—2019年度花后35—45 d连续10 d降雨,该年度小麦穗发芽严重。

图1

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图12018和2019年小麦花后日降雨量及累积降雨量

Fig. 1Daily and cumulative rainfall after flowering of wheat in 2018 and 2019

1.2 测定项目及方法

1.2.1 田间自然穗发芽率的测定 参照朱冬梅等^[21]的方法,小麦成熟后在田间自然淋雨3 d以上,直至对照穗发芽时,取样调查穗发芽率。本试验第1年小麦正常收获期无雨,小区收获一半,留一半于田间延迟收获,直至后面3 d连续下雨对照穗发芽,随即取样调查发芽率;第2年由于收获期一直降雨,收获时已有穗发芽发生,则直接取样调查发芽率。取样时各处理分别取15穗,剥粒调查发芽情况,以露白为最低发芽标准。田间自然穗发芽率（%）=发芽籽粒数/15穗籽粒总数×100%

1.2.2 穗发芽率及粒发芽率测定 在生理成熟期和蜡熟期各处理分别取10个穗子,其中5个穗子用于测定穗发芽,5个穗子用于测定粒发芽。穗发芽参照马文洁等^[22]的方法加以改进,即先将穗子用3%的NaClO表面消毒30 min,再用无菌水冲洗干净,然后将穗子插在试管中置于25℃恒温光照培养箱中培养,其间采用喷雾方式补充适量蒸馏水以保持穗子湿度,7 d后取出剥粒记录发芽种子数（以种子露白为最低发芽标准）,计算穗发芽率（%）=发芽籽粒数/穗籽粒总数×100%。粒发芽参照原亚萍等^[2]的方法加以改进,即将穗子手工脱粒后,随机数出100粒,用1%的NaClO溶液浸泡10 min,再用无菌水冲洗干净,然后将籽粒腹沟朝下摆放在铺有两层滤纸且经过高温消毒的培养皿中,用3 mL蒸馏水浸湿滤纸,最后将培养皿放置在20℃的恒温光照培养箱中暗化发芽,每天喷雾适量蒸馏水,7 d后统计发芽率,以芽长达到种子长的一半时为最低发芽标准,粒发芽率（%）=发芽数/总粒数×100%。生理成熟期以小麦穗颈和颖壳由绿转黄为判断标准^[23],本试验中供试小麦品种经过田间观察,在开花后35 d即进入这一时期,故生理成熟期时间定为花后35 d。

1.2.3 α-淀粉酶活性测定 开花后15、20、25、30、35、40、45 d,分别取每个处理4—5个穗子的中部籽粒,置于液氮中保存,然后转移到-80℃冰柜中放置待测。所用生化试剂购于Sigma公司,试剂盒由北京索莱宝科技有限公司提供,应用双抗体夹心酶联免疫吸附法（Elisa）测定。

1.2.4 淀粉及可溶性糖含量动态测定 取样时间同1.2.3,将所取4—5个穗子的中部籽粒置于105℃烘箱中杀青10—30 min,然后75℃烘干至恒重。采用蒽酮比色法^[24]测定淀粉和可溶性糖含量。

1.2.5 小麦籽粒及面粉品质测定 将收获的籽粒后熟2个月后磨粉,测定籽粒蛋白质含量^[25]、湿面筋含量^[26]、沉降值、降落值^[27]和面粉RVA特征值^[28]。

1.2.6 面粉中总淀粉、直链淀粉及支链淀粉的测定 样品来源同1.2.5,总淀粉含量采用蒽酮比色法^[24]测定,直链淀粉按照GB/T 15683-1995测定^[29],总淀粉含量减去直链淀粉含量为支链淀粉含量。

1.3 统计分析

所有试验数据使用Microsoft Excel 2013进行汇总;用DPS 7.05系统软件进行统计分析,采用LSD法进行显著性比较分析;用Origin 2018进行作图。

2 结果

2.1 外源ABA喷施时期和喷施浓度对冬小麦穗发芽与粒发芽性状的影响

2个小麦试验生长季两粒色品种、不同喷施时期与ABA浓度均显著影响田间自然穗发芽率、生理成熟期及蜡熟期的穗、粒发芽率;同时,品种（A）×浓度（C）交互作用几乎对所有指标差异均显著（2018年蜡熟期的粒发芽率除外）,但整体以品种、喷施时期、ABA浓度三个因素的主效应更大（表1）。进一步分析表明（表2）,品种间中科麦138发芽率显著高于绵麦367（蜡熟期粒发芽率差异不显著）,2018年和2019年绵麦367田间自然穗发芽率分别较中科麦138低43.0和46.5个百分点。不同时期喷施ABA对小麦发芽均有抑制作用,2个小麦试验生长季均以花后30 d（30 DAA）喷施抑制发芽效果更好,（15+30）DAA处理下田间自然穗发芽率、生理成熟期和蜡熟期的穗发芽率和粒发芽率反而更高;从喷施浓度处理看,2018年以喷施浓度C₁处理效果好,中科麦138生理成熟期及蜡熟期的粒发芽率较CK下降13.8和3.8个百分点,而绵麦367的生理成熟期及蜡熟期的粒发芽率较CK下降23.5和9.7个百分点;2019年以喷施浓度C₂处理效果好,中科麦138的生理成熟期及蜡熟期粒发芽率较CK下降22.5和19.6个百分点,绵麦367较CK下降10.0和12.0个百分点,说明多雨年份可降低品种对ABA的敏感性,喷施浓度需要提高。

Table 1
表1
表1喷施时期和ABA浓度对两粒色小麦品种穗、粒发芽性状的方差分析
Table 1ANOVA of ABA concentration and spraying period on spike and grain germination traits of white and red wheat cultivar

处理 Treatment		田间自然穗发芽率 Spike germination rate in field	生理成熟期 Physiological maturity (35DAA)		蜡熟期 Dough stage (45DAA)
			穗发芽率 Spike germination rate	粒发芽率 Grain germination rate	穗发芽率 Spike germination rate	粒发芽率 Grain germination rate
2018	品种Cultivar (A)	15860.1**	29.5*	524.3**	650.8**	1.2ns
	喷施时期Spraying period (B)	69.4**	222.6**	219.6**	543.9**	0.7ns
	ABA浓度ABA concentration (C)	371.5**	145.8**	234.0**	59.0**	15.9**
	品种×喷施时期 A×B	4.5ns	63.6**	22.8**	0.0ns	2.4ns
	品种×ABA浓度 A×C	36.0**	5.0*	36.4**	4.4*	1.9ns
	喷施时期×ABA浓度 B×C	1.7ns	0.4ns	3.6ns	8.6**	3.1ns
	品种×喷施时期×ABA浓度A×B×C	3.8*	0.4ns	9.8**	5.2*	5.0*
2019	品种Cultivar (A)	1774.4**	921.0**	33.1*	200.5**	1.0ns
	喷施时期Spraying period (B)	8.2*	99.7**	6.6*	21.9**	16.8**
	ABA浓度ABA concentration (C)	306.4**	483.0**	96.4**	154.1**	40.8**
	品种×喷施时期 A×B	4.7*	6.2*	2.3ns	3.3ns	0.2ns
	品种×ABA浓度 A×C	14.8**	11.9**	14.8**	6.2**	5.9**
	喷施时期×ABA浓度 B×C	9.3**	3.4*	2.7ns	4.8**	2.5ns
	品种×喷施时期×ABA浓度A×B×C	9.0**	2.5ns	1.5ns	1.0ns	3.4*

*和**分别表示0.05和0.01的显著水平,ns表示差异不显著。发芽率数据均经反正弦转换后进行分析。下同
* and ** represented significance at 0.05 and 0.01 levels, and ns represented no significance. The data of germination rate were analyzed by inverse sine conversion. The same as below

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Table 2
表2
表2喷施时期和ABA浓度对两粒色小麦品种穗发芽与粒发芽的影响
Table 2Effects of ABA concentration and spraying period on spike germination and grain germination of white and red wheat cultivar

年份 Year	处理 Treatment		田间自然穗发芽率 Spike germination rate in field (%)		生理成熟期Physiological maturity (35DAA)				蜡熟期Dough stage (45DAA)
					穗发芽率 Spike germination rate (%)		粒发芽率 Grain germination rate (%)		穗发芽率 Spike germination rate (%)		粒发芽率 Grain germination rate (%)
			ZKM138	MM367	ZKM138	MM367	ZKM138	MM367	ZKM138	MM367	ZKM138	MM367
2018	喷施时期 Spraying period	B1	45.4a	2.9b	11.3b	10.6b	32.9b	12.7b	72.4b	62.0b	91.7a	91.0a
		B2	47.3a	3.9a	17.5a	12.4a	51.2a	19.3a	82.9a	72.5a	96.2a	91.3a
	ABA浓度 ABA concentration	CK	51.9a	7.2a	22.7a	16.7a	49.8a	30.0a	85.1a	81.2a	95.3a	95.2a
		C1	42.8b	0.9c	8.8c	7.2c	36.0c	6.5c	70.0c	53.2c	91.5b	85.5b
		C2	44.4b	2.1b	11.7b	10.7b	40.3b	11.5b	77.9b	67.3b	95.0a	92.8a
		平均Mean	46.4a	3.4b	14.4a	11.5b	42.1a	16.0b	77.6a	67.3b	93.9a	91.2a
2019	喷施时期 Spraying period	B3	65.7a	17.1b	29.6a	19.7b	38.6ab	29.9a	76.0a	66.6a	83.6a	82.9a
		B1	65.6a	16.8b	23.2b	17.3c	34.4b	28.7a	67.1b	62.1b	73.0b	73.2b
		B4	66.5a	24.2a	30.4a	22.0a	43.8a	30.9a	79.7a	67.8a	87.7a	85.7a
	ABA浓度 ABA concentration	CK	81.3a	32.4a	38.6a	28.6a	49.1a	35.0a	87.2a	75.0a	92.2a	85.9a
		C1	67.6b	14.8b	27.9b	17.5b	41.1b	29.4b	72.3b	65.7b	79.4b	82.0a
		C2	48.9c	11.0c	16.7c	12.8c	26.6c	25.0c	63.3c	55.8c	72.6c	73.9b
		平均Mean	65.9a	19.4b	27.7a	19.7b	38.9a	29.8b	74.3a	65.5b	81.4a	80.6a

DAA表示花后天数;B₁：花后30 d;B₂：花后35 d;B₃：花后15 d;B₄：花后15 d和30 d叠加;CK：清水;C₁：50 mg·L^-1ABA;C₂：100 mg·L^-1ABA。各处理间同一列或平均值间同一行的小写字母a、b、c表示 0.05 水平差异显著。发芽率数据均经反正弦转换后进行分析。下同
DAA represented the days after anthesis; B₁: 30 days after anthesis; B₂: 35 days after anthesis; B₃: 15 days after anthesis; B₄:15 days after anthesis plus 30 days; CK: Water; C₁: 50 mg·L^-1ABA; C₂: 100 mg·L^-1ABA. Values followed by different letters within a column indicated significantly different at 0.05 level. The data of germination rate were analyzed by inverse sine transformation. The same as below

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2.2 外源ABA对冬小麦籽粒α-淀粉酶活性及淀粉和可溶性糖含量影响

2.2.1 α-淀粉酶活性 冬小麦花后15—45 d,籽粒α-淀粉酶活性变化总体呈降升交替的“W”型变化趋势（图2）。α-淀粉酶的活性在花后35 d（生理成熟期）降至最低,而在花后45 d（蜡熟期）又升高。花后45 d白皮易穗发芽品种中科麦138的α-淀粉酶活性较红皮抗穗发芽品种绵麦367提高25.8%,但不同喷施时期处理间α-淀粉酶活性的差异不显著,而随ABA浓度的增加,α-淀粉酶活性呈显著降低趋势。花后30 d喷施浓度C₂处理对α-淀粉酶活性的抑制作用最强,花后45 d中科麦138和绵麦367的α-淀粉酶活性分别较CK降低28.0%和32.8%。

图2

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图2喷施时期和ABA浓度对籽粒α-淀粉酶活性的影响

CK：清水;C₁：50 mg·L^-1ABA;C₂：100 mg·L^-1ABA。图中三线表为花后45 d α-淀粉酶活性的方差分析,F_C：品种间的F值,F_SP：喷药时期间的F值,F_ABAC：脱落酸浓度间的F值。^**表示0.01水平差异显著性。下同
Fig. 2Effects of ABA concentration and spraying period on dynamic changes of amylase activity in wheat

CK: Water; C₁: 50 mg·L^-1ABA; C₂: 100 mg·L^-1ABA. The three-line table in the figure was the analysis of variance of α-amylase activity 45 days after anthesis. F_C: F value between cultivar, F_SP: F value between spraying period, F_ABAC: F value between ABA concentration. ^** indicated significant difference at 0.01 level. The same as below


2.2.2 淀粉和可溶性糖含量 从灌浆期淀粉和可溶性糖含量变化来看,两者为此起彼落的关系（图3）。灌浆期淀粉含量在花后20 d迅速上升,花后35—40 d达最大峰值,而在花后40 d或45 d下降,对应于此期的可溶性糖含量有所上升,预示着淀粉的降解。花后40 d和45 d中科麦138的可溶性糖含量分别较绵麦367提高34.7%和59.7%。喷施ABA处理均可促进可溶性糖转化为淀粉,总淀粉含量增加,且随喷施浓度增加而增加。在淀粉含量达到最大值时,C₁浓度下,中科麦138和绵麦367籽粒淀粉含量分别较CK提高4.3和4.6个百分点,C₂浓度下分别较CK提高7.0和6.7个百分点。但过早喷施（15 DAA处理）和叠加喷施（（15+30）DAA处理）均使中科麦138淀粉降解早于花后30 d喷施处理,这可能是过早或叠加喷施处理加快籽粒成熟的原因。并且,花后30 d喷施50—100 mg·L^-1ABA处理可显著延缓两粒色品种淀粉的降解,且以C₂浓度延缓作用最强,该浓度处理下中科麦138花后45 d的淀粉含量较花后40 d下降2.5个百分点,同期CK下降4.0个百分点,而绵麦367花后45 d的淀粉含量亦较花后40 d下降5.9个百分点,同期CK下降12.8个百分点。

图3

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图3喷施时期和ABA浓度对小麦淀粉及可溶性糖含量的影响

柱状图为总淀粉含量,折线图为可溶性糖含量
Fig. 3Effects of ABA concentration and spraying period on dynamic changes of total starch and soluble sugar in wheat

The column showed the total starch content and the broken line shows the soluble sugar content

2.3 外源ABA对穗发芽抑制效果的解除

在收获后30 d和60 d测定籽粒发芽率,结果表明不同时期中科麦138的籽粒发芽率均低于绵麦367,但ABA的抑制萌发效应在2个品种间表现略有不同（图4）。在30 DAA和（15+30）DAA喷施ABA对中科麦138收获后30 d的籽粒发芽仍存在一定抑制作用,其籽粒发芽率显著低于CK,但对收获后60 d的籽粒发芽率却有显著提升作用,发芽率均在98%以上,15 DAA喷施ABA的籽粒在收获后30 d和60 d的发芽率与CK无显著差异;而对绵麦367而言,各时期喷施ABA对其收获后30 d和60 d的籽粒发芽率均无不良影响,发芽率均在93%以上。由此可以表明,花后30 d喷施50—100 mg·L^-1ABA可有效抑制穗发芽,其抑制效应均在收获后60 d全部解除,不影响第2年作为种用的籽粒发芽率。

图4

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图4喷施时期和ABA浓度对小麦籽粒发芽抑制效应解除时间

柱状图上不同小写字母表示0.05水平异显著。DAH：收获后天数;DAA：花后天数
Fig. 4Effects of ABA concentration and spraying period on release time of seed germination inhibition effect in wheat

Values followed by different letters within a bar chart indicated significantly different at 0.05 level. DAH: Days after harvest; DAA: Days after anthesis

2.4 外源ABA对小麦籽粒和面粉品质的影响

2.4.1 蛋白质品质和淀粉品质 相比蛋白质含量和质量,降落值、峰值粘度、崩解值等淀粉品质性状受试验因子的影响更大（表3）。2个小麦试验生长季品种间蛋白质含量差异不显著,而沉淀值、降落值、峰值粘度、崩解值差异显著;喷施时期在2018年显著影响蛋白质含量、降落值和峰值粘度,2019年则对所有指标均有显著影响;2个小麦试验生长季,喷施浓度均显著影响沉淀值、降落值、峰值粘度和崩解值,2019年还显著影响了湿面筋含量。品种（A）×ABA浓度（C）的互作效应除2018年粗蛋白含量和2019年湿面筋含量外,对其他指标均有显著影响;喷施时期（B）×ABA浓度（C）的互作效应2018年显著影响粗蛋白含量、沉淀值和降落值,2019年则对所有指标均有显著影响,但品种、喷施ABA时期和浓度三者间互作效应的影响相对占比较小。

Table 3
表3
表3喷施时期和ABA浓度对蛋白质和淀粉品质的方差分析
Table 3ANOVA of ABA concentration and spraying period on protein and starch quality of wheat

年份 Year	处理 Treatment	粗蛋白含量 Protein content	沉淀值 Sedimentation value	湿面筋含量 Wet gluten	降落值 Falling number	峰值粘度 Peak viscosity	崩解值 Breakdown value
2018	品种Cultivar (A)	0.0ns	243.6**	521.1**	6348.7**	6900.5**	1313.9**
	喷施时期Spraying period (B)	7.8*	3.8ns	3.4ns	116.0**	8.5*	1.3ns
	ABA浓度ABA concentration (C)	0.3ns	8.7**	0.1ns	174.9**	81.1**	15.9**
	品种×喷施时期 A×B	2.2ns	1.2ns	3.4ns	98.0**	12.3*	4.1ns
	品种×ABA浓度 A×C	0.1ns	10.7**	14.1**	5.5*	8.0**	14.4**
	喷施时期×ABA浓度 B×C	5.0*	20.8**	2.1ns	11.6**	0.3ns	1.0ns
	品种×喷施时期×ABA浓度A×B×C	1.0ns	17.4**	9.0**	0.9ns	1.8ns	0.5ns
2019	品种Cultivar (A)	7.1ns	108.5**	2.1ns	21952.0**	1116.8**	1249.7**
	喷施时期Spraying period (B)	7.6*	9.2**	5.9*	44.1**	96.8**	125.3**
	ABA浓度ABA concentration (C)	1.7ns	26.1**	62.2**	68.3**	91.6**	80.3**
	品种×喷施时期 A×B	1.2ns	8.1*	29.3**	18.0**	10.5**	16.3**
	品种×ABA浓度 A×C	3.4*	20.8**	1.0ns	38.8**	6.7**	4.4*
	喷施时期×ABA浓度 B×C	7.2**	13.9**	12.1**	4.7**	6.2**	5.7**
	品种×喷施时期×ABA浓度A×B×C	3.8*	17.1**	0.5ns	6.1**	5.4**	5.5**

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进一步分析表明（表4）,品种间主效应表现为2018年中科麦138沉淀值、降落值、峰值粘度和崩解值显著高于绵麦367,2019年则与之相反,以绵麦367更高;湿面筋含量2个小麦试验生长季均以中科麦138更高,仅2018年存在显著差异。除粗蛋白含量外,2个小麦试验生长季均以B₁喷施处理的（30 DAA）沉淀值和湿面筋含量更高,而降落值、峰值粘度和崩解值在2018年以B₁喷施处理（30DAA）更高,2019年则以B₃喷施处理（15DAA）更高,分别较B₁处理提高16.7%、18.5%和15.5%。与CK相比,喷施ABA后均能提高沉淀值、降落值、峰值粘度和崩解值,且呈随喷施浓度的增大而增加的趋势,2个小麦试验生长季均以喷施浓度C₂效果更好,其沉淀值、降落值、峰值粘度和崩解值分别较CK提高4.3%—8.8%、20.9%—24.2%、26.5%—51.4%和12.4%—43.4%。

Table 4
表4
表4喷施时期和ABA浓度对蛋白质和淀粉品质的影响
Table 4Effects of ABA concentration and spraying period on protein and starch quality of wheat

年份 Year	处理 Treatment		粗蛋白含量 Protein content (%)	沉淀值 Sedimentation value (mL)	湿面筋含量 Wet gluten (%)	降落值 Falling number (s)	峰值粘度 Peak viscosity (RVU)	崩解值 Breakdown value (RVU)
2018	品种 Cultivar	ZKM138	8.0a	13.5a	10.6a	281a	137.2a	95.8a
		MM367	8.0a	10.2b	5.8b	168b	83.9b	69.5b
	喷施时期 Spraying period	B1	8.1a	12.1a	8.3a	232a	114.1a	83.7a
		B2	7.8b	11.6a	8.1a	219b	107.0b	81.6a
	ABA浓度 ABA concentration	CK	8.0a	11.7b	8.2a	198c	96.6c	77.6c
		C1	8.0a	11.7b	8.2a	233b	112.9b	83.3b
		C2	8.0a	12.2a	8.2a	246a	122.2a	87.2a
		平均Mean	8.0	11.9	8.2	225	110.6	82.7
2019	品种 Cultivar	ZKM138	9.2a	13.0b	13.3a	65b	11.1b	13.7b
		MM367	9.1a	17.2a	13.2a	123a	32.4a	34.0a
	喷施时期 Spraying period	B3	9.1b	15.1ab	12.8b	105a	25.0a	26.9a
		B1	9.0b	15.8a	13.5a	90b	21.1b	23.3b
		B4	9.4a	14.5b	13.4 a	87b	19.1c	21.4c
	ABA浓度 ABA concentration	CK	9.2a	14.8b	12.1c	86c	17.3c	19.6c
		C1	9.1a	14.4b	13.3b	91b	21.8b	23.9b
		C2	9.2a	16.1a	14.4a	104a	26.2a	28.1a
		平均Mean	9.2	15.1	13.3	94	21.8	23.9

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2.4.2 淀粉含量及组分 中科麦138的总淀粉含量、支链淀粉含量较绵麦367均显著提高,其平均增幅为17.9%、28.5%,但其直链淀粉含量、直/支比显著降低,平均降幅分别为13.2%、33.3%（表5）。B₁处理（30 DAA）和B₃（15 DAA）处理通过增加支链淀粉含量进而增加总淀粉含量,而叠加施用B₄（15+30）DAA处理则会降低支链淀粉含量进而减少总淀粉含量。随喷施浓度增加,支链淀粉含量增加,进而总淀粉含量增加,喷施ABA浓度C₁、C₂处理后支链淀粉含量分别较CK增加4.4和8.1个百分点,总淀粉含量增加3.9和7.6个百分点,直/支比下降9.1%和18.2%。

Table 5
表5
表5施药时期和ABA浓度对面粉总淀粉及其组分含量的影响
Table 5Effects of ABA concentration and spraying period on the contents of total starch and its components in wheat flour

处理 Treatment		直链淀粉含量 Amylase content (%)	支链淀粉含量Amylopectin content (%)	直/支比 Amylase/Amylopectin ratio	总淀粉含量 Starch content (%)
品种 Cultivars	ZKM138	13.2b	54.6a	0.24b	67.8a
	MM367	15.2a	42.5b	0.36a	57.5b
喷施时期 Spraying period	B3	14.2a	48.9a	0.30b	63.0a
	B1	14.1a	50.7a	0.29c	64.6a
	B4	14.2a	46.1b	0.32a	60.4b
ABA浓度 ABA concentration	CK	14.4a	44.4c	0.33a	58.8c
	C1	14.2b	48.8b	0.30b	62.7b
	C2	13.9c	52.5a	0.27c	66.4a
F值 F-value	品种Cultivar (A)	1080.0**	191.3**	157.8**	129.8**
	喷施时期Spraying period (B)	1.8ns	15.6**	27.1**	13.7**
	ABA浓度ABA concentration (C)	27.4**	65.0**	79.5**	51.3**
	品种×喷施时期 A×B	9.0**	1.9ns	0.4ns	2.5ns
	品种×ABA浓度 A×C	4.3*	4.6*	1.7ns	5.1*
	喷施时期×ABA浓度 B×C	1.9ns	2.4ns	5.4**	2.0ns
	品种×喷施时期×ABA浓度A×B×C	1.6ns	3.2*	2.7ns	2.6ns

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3 讨论

3.1 外源喷施ABA对α-淀粉酶活性的影响

已有研究表明,种子的萌发与休眠受激素调控,ABA促进休眠,而GA促进萌发^[11],因而施用外源ABA抑制穗发芽有坚实的理论基础。但何时施用、施用何种浓度值得考究。关于种子发育过程中ABA的变化,有研究认为随种子发育进程推进,ABA含量不断增加,又随籽粒成熟而下降^[30],并认为ABA是促进籽粒脱水成熟的重要因素^[31],同时,小麦品种对ABA的敏感性具有随胚萌发能力的提高而下降的特点^[32]。关于外源ABA应用技术及调控对象也有多个方面,如早期喷施可用于籽粒灌浆的调控^[19],晚期喷施可促进脱水,加快种子成熟^[31],也可用于穗发芽的调控^[33]。因而根据调控目的,ABA的施用时期和施用浓度非常重要。雍太文等^[13]在水稻上研究发现,ABA喷期越迟,抑制穗发芽效果越好,且迟喷（灌浆中期或乳熟末期）50 mg·L^-1ABA处理穗发芽率最低,而早喷（授粉末期）则需要适当增加喷施浓度,以90 mg·L^-1ABA处理穗发芽率最低,对籽粒的抑制作用均可在收获后30 d解除。本试验基于田间自然穗发芽及室内受控条件下的穗发芽与粒发芽调查结果,表明花后30 d喷施ABA对小麦穗、粒发芽的抑制效果最优,收获期雨水较少年份50 mg·L^-1处理即可,收获期雨水较多年份100 mg·L^-1效果较好。与此同时,本研究还发现,ABA对籽粒发芽的抑制作用于收获后60 d全部解除,不会影响第2年作为种用的正常发芽率。

α-淀粉酶作为淀粉分解的关键酶,其活性大小是鉴定小麦穗发芽的主要指标^[34]。前人研究认为ABA、SA等穗发芽抑制剂对α-淀粉酶活性均有显著抑制作用^{[33, 35-36]},小麦穗发芽率与籽粒可溶性糖含量及α-淀粉酶活性呈正相关关系^[37]。本研究结果也表明,α-淀粉酶活性在种子发育过程中呈现“W”型变化趋势,2个低谷分别出现在开花后20 d和开花后35 d,以第2个低谷谷底更低,也预示着种子生理成熟,具备发芽能力。籽粒发育过程中淀粉和可溶性糖含量也存在此起彼落的关系。α-淀粉酶在籽粒发育后期主要促进淀粉水解为可溶性糖,从而为籽粒萌发提供能量,因而在穗发芽中其后期的活性备受关注。田间调查表明,2个品种花后35 d即进入生理成熟期。前人研究认为^[23],生理成熟期种子即具备发芽的能力,只要条件适宜,即可穗上发芽。因而,本文研究认为,花后35—45 d是穗发芽敏感期,该阶段籽粒a-淀粉酶活性高低与穗发芽关系密切。本研究也发现,白粒的中科麦138花后35 d的α-淀粉酶活性高于红粒的绵麦367,可能也是绵麦367穗发芽低于中科麦138的内在生理原因。花后不同时期喷施ABA后均可抑制花后35—45 d的α-淀粉酶活性,以花后30 d喷施抑制作用最强,而花后15 d喷施和花后（15+30）d叠加施用会导致淀粉降解提前,可能与过早喷施促进早熟有关^[38],同时,小麦品种对ABA的敏感性具有随胚萌发能力的提高而下降的变化特点^[32],这也可能是花后35 d喷施抑制穗发芽效果甚微的原因。就施用浓度而言,随施用浓度增加,抑制作用增强,淀粉水解减缓。花后30 d喷施100 mg·L^-1 ABA处理下,中科麦138花后45 d的淀粉含量较花后40 d下降2.5个百分点,同期CK下降4.0个百分点;而绵麦367花后45 d的淀粉含量亦较花后40 d下降5.9个百分点,同期CK下降12.8个百分点。可见,针对穗发芽而言,花后30 d喷施50—100 mg·L^-1 ABA,可有效抑制α-淀粉酶的转录与其酶自身的活性,抑制穗发芽敏感期淀粉的水解,达到抑制穗发芽的目的。

3.2 外源喷施ABA对淀粉组分含量及糊化特性的影响

前人有关ABA信号转导与代谢调控进而影响籽粒蛋白质品质的研究较多,如赵虎成等^[39]发现,ABA能诱导许多基因的表达,从而调控蛋白质的合成;崔志青等^[20]研究结果表明,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。本研究亦证实ABA对蛋白质品质具有调控效应,以花后30 d喷施（30 DAA）处理的沉淀值和湿面筋含量更高,且有随ABA浓度增加呈增大趋势,100 mg·L^-1ABA处理的沉淀值较CK提高4.3%—8.8%。

相比蛋白质品质而言,本研究认为ABA更多地影响了籽粒淀粉组分含量及面粉糊化特性。本研究结果表明,不同ABA浓度处理后支链淀粉含量和总淀粉含量提高,直/支比下降。直链淀粉和支链淀粉作为小麦籽粒淀粉的组成成分,其含量和比例是造成不同小麦品种面粉糊化特性差异的决定因素^[40]。前人研究结果表明,降低直链淀粉或增加支链淀粉的含量,降低直/支比,可提高小麦粉的膨胀力,降低糊化凝胶硬度,增强糊化淀粉粒的形变能力,促使面条质地软而富有弹性^[41,42]。小麦峰值粘度和崩解值与支链淀粉含量呈极显著正相关,而与直链淀粉含量及直/支比呈显著负相关^[43],高的支链淀粉含量具有较高的粘度^[44]。本研究结果表明,ABA喷施后面粉糊化特性改善,100 mg·L^-1浓度的ABA处理的降落值提高20.9%—24.2%,峰值粘度提升26.5%—51.4%,崩解值提升12.4%—43.4%。2018年以花后30 d喷施处理的降落值、峰值粘度和崩解值更高,2019年则以花后15 d喷施处理更高。而有关ABA调控淀粉合成与累积的机理,前人认为可能与外源ABA喷施后提高了籽粒ABA水平及ABA与GAs比值,增强籽粒中蔗糖-淀粉代谢途径关键酶活性及相关蛋白质表达,从而促进籽粒淀粉合成和累积有关^[45,46,47],同时,淀粉组分与淀粉的粒度分布密切相关,A型淀粉粒体积比例与淀粉的直/支比呈显著正相关关系,B型淀粉粒体积比例与淀粉的直/支比呈显著负相关关系^[48]。本研究发现ABA不仅可抑制穗发芽,还对淀粉质量提升有重要作用,但有关ABA喷施后促进支链淀粉含量增加,进而增加总淀粉含量优化糊化特性的机制还需要进一步深入研究。

4 结论

在花后30 d喷施50—100 mg·L^-1ABA,可有效降低穗发芽敏感期（花后35—45d）α-淀粉酶活性,延缓生理成熟后淀粉的水解,减少萌发能量物质可溶性糖的供给,从而达到抑制小麦穗发芽的目的。喷施后还可增加支链淀粉含量进而增加总淀粉含量、降低直/支比,改善面粉糊化特性,并对湿面筋含量和沉降值也有一定提升作用,切实起到有灾减灾、无灾提质的作用。

（责任编辑 杨鑫浩）

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Journal of Triticeae Crops, 2000,20(2):93-95. (in Chinese)
DOI:10.7606/j.issn.1009-1041.2000.02.100URL [本文引用: 1]
Based on the results of many years research of preharvest sprouting physiological mechanism both at laboratory and field, a preharvest sprouting inhibiting agent(N 5) was invented. It was found that the best spraying period was 36 days after flowering. W

[15] 张玲娥, 张萍 . 植物生长调节剂对冬小麦籽粒灌浆期源库关系的调控
核农学报, 2005,19(3):228-231.
URL [本文引用: 1]
Taking relationship, translocation and distribution of photosynthesis assimilates as indexes, the adjustment of plant growth regulator in source-sink was investigated by ~(14)C-tracer technique during grain filling of winter wheat. The results show that the application of ABA and GA_3 affect the translocation and distribution of the ~(14)C-assimilates. Application of ABA inhibits and GA_3 stimulats effect on the translocation and distribution of ~(14)C-assimilates.
ZHANG L E, ZHANG P . Adjustment of plant growth regulations in source-sink relationship during grain filling of winter wheat
Acta Agriculturae Nucleatae Sinica, 2005,19(3):228-231. (in Chinese)
URL [本文引用: 1]
Taking relationship, translocation and distribution of photosynthesis assimilates as indexes, the adjustment of plant growth regulator in source-sink was investigated by ~(14)C-tracer technique during grain filling of winter wheat. The results show that the application of ABA and GA_3 affect the translocation and distribution of the ~(14)C-assimilates. Application of ABA inhibits and GA_3 stimulats effect on the translocation and distribution of ~(14)C-assimilates.

[16] 李文阳, 尹燕枰, 闫素辉, 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.
DOI:10.3321/j.issn:1000-7091.2007.01.002URL [本文引用: 1]
选用2个粒重差异较大的小麦品种济麦20(小粒型)和山农710331(大粒型)，对比研究其籽粒发育过程中4种主要内源激素(IAA，GAs，ZR和ABA)的含量变化及籽粒灌浆特征。结果表明，2个品种内源激素含量变化动态趋势基本一致，但品种间内源激素含量存在基因型差异。在籽粒发育过程中，IAA，GAs和ZR含量呈籽粒发育前期高、后期低的趋势，而ABA含量呈&ldquo;V&rdquo;型曲线变化。用Logistic方程拟合籽粒灌浆过程可以看出，籽粒灌浆启动时间、灌浆速率以及灌浆持续时间共同决定小麦最终籽粒重的高低。与小粒型品种济麦20相比，大粒型品种山农710331籽粒发育初期较高的ZR含量使其籽粒灌浆启动时间早；在整个籽粒发育过程中较高的GAs，ZR和ABA含量与较高的籽粒灌浆速率相联系；籽粒发育后期较高的ZR含量及相对较低的ABA增幅，可能是其籽粒灌浆持续时间较长的一个重要原因。
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
Journal of North China Agronomy, 2007, 22(1):5-8. (in Chinese)
DOI:10.3321/j.issn:1000-7091.2007.01.002URL [本文引用: 1]
选用2个粒重差异较大的小麦品种济麦20(小粒型)和山农710331(大粒型)，对比研究其籽粒发育过程中4种主要内源激素(IAA，GAs，ZR和ABA)的含量变化及籽粒灌浆特征。结果表明，2个品种内源激素含量变化动态趋势基本一致，但品种间内源激素含量存在基因型差异。在籽粒发育过程中，IAA，GAs和ZR含量呈籽粒发育前期高、后期低的趋势，而ABA含量呈&ldquo;V&rdquo;型曲线变化。用Logistic方程拟合籽粒灌浆过程可以看出，籽粒灌浆启动时间、灌浆速率以及灌浆持续时间共同决定小麦最终籽粒重的高低。与小粒型品种济麦20相比，大粒型品种山农710331籽粒发育初期较高的ZR含量使其籽粒灌浆启动时间早；在整个籽粒发育过程中较高的GAs，ZR和ABA含量与较高的籽粒灌浆速率相联系；籽粒发育后期较高的ZR含量及相对较低的ABA增幅，可能是其籽粒灌浆持续时间较长的一个重要原因。

[17] 戴忠民 . 喷施6BA和ABA对冬小麦籽粒胚乳细胞增殖和淀粉积累的影响
麦类作物学报, 2008,28(3):134-139.
[本文引用: 2]

DAI Z M . Effects of 6BA and ABA on endosperm cell propagation and starch accumulation in grains of winter wheat
Journal of Triticeae Crops, 2008,28(3):134-139. (in Chinese)
[本文引用: 2]

[18] 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(5):899-906.
DOI:10.3724/SP.J.1006.2009.00899URL [本文引用: 1]
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.
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 Agronomica Sinica, 2009,35(5):899-906. (in Chinese)
DOI:10.3724/SP.J.1006.2009.00899URL [本文引用: 1]
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.

[19] YANG J C, ZHANG J H, WANG Z Q, ZHU Q S . Hormones in the grains in relation to sink strength and postanthesis development of spikelets in rice
Plant Growth Regulation, 2003,41(3):185-195.
DOI:10.1023/B:GROW.0000007503.95391.38URL [本文引用: 2]
Inferior spikelets usually exhibit a slower grain filling rate and lower grain weight than superior spikelets in a rice (Oryza sativa L.) panicle. This study investigated whether the variations in grain filling between the two kinds of spikelets were attributed to their sink strength and whether the sink strength was regulated by the hormonal levels in the grains. Using two field-grown rice genotypes, the division rate of endosperm cells, hormonal levels in the grains, and grain weight of both superior and inferior spikelets were determined during the grain filling period. The results showed that superior spikelets had dominance over inferior spikelets in endosperm cell division rate and cell number, grain filling and grain weight. Changes in zeatin (Z) and zeatin riboside (ZR) contents paralleled and were very significantly correlated with the cell division rate and cell number. Cell division rate and the content of indole-3-acetic acid (IAA) in the grains were also significantly correlated. Gibberellin (GAs; GA₁+ GA₄) content of the grains was high but ABA levels were low at the early grain filling stage. ABA increased substantially during the linear phase of grain growth and was very significantly correlated with grain dry weight during this period. Application of kinetin at 2 through 6 days post anthesis (DPA) significantly increased cell number, while spraying ABA at 11 through 15 DPA significantly increased the grain filling rate. The results suggest that differences in sink strength are responsible for variations in grain filling between superior and inferior spikelets. Both cytokinins and IAA in the grains may mediate cell division in rice endosperm at early grain filling stages, and therefore regulate the sink size of the grain, whereas ABA content correlates with sink activity during the linear period of grain growth.

[20] 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 . 喷施ABA对小麦籽粒谷蛋白组分含量及GMP粒度分布的影响
中国农业科学, 2010,43(12):2595-2602.
URL [本文引用: 2]
【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&#8226;kg-1, 45 mL&#8226;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.


CUI Z Q, HE D X, CAI T, WANG C Y, WANG G C, MENG F Y, HAN Z H, LI N N, WANG Z L . Effects of spraying ABA on glutenin fraction content and GMP size distribution in wheat grain
Scientia Agricultura Sinica, 2010,43(12):2595-2602. (in Chinese)
URL [本文引用: 2]
【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&#8226;kg-1, 45 mL&#8226;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.


[21] 朱冬梅, 李曼, 李东升, 吴素兰, 张晓, 张伯桥 . 长江中下游麦区小麦新品种穗发芽抗性及鉴定方法研究
核农学报, 2018,32(4):795-801.
[本文引用: 1]

ZHU D M, LI M, LI D S, WU S L, ZHANG X, ZHANG B Q . Evaluation and determination method of pre-harvest sprouting resistance of new wheat cultivars in middle and lower reaches of Yangtze River
Journal of Nuclear Agricultural Sciences, 2018,32(4):795-801. (in Chinese)
[本文引用: 1]

[22] 马文洁, 张传量, 宋晓朋, 冯洁, 崔紫霞, 孙道杰 . 不同麦区小麦品种穗发芽抗性及其与穗部性状的相关性
麦类作物学报, 2016,36(10):1269-1274.
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MA W J, ZHANG C L, SONG X P, FENG J, CUI Z X, SUN D J . Pre-harvest sprouting resistance in wheat from different wheat regions and its correlation with ear characteristics
Journal of Triticeae Crops, 2016,36(10):1269-1274. (in Chinese)
[本文引用: 1]

[23] 肖世和, 闫长生, 张海萍, 孙果忠 . 小麦穗发芽研究. 北京: 中国农业科学技术出版社, 2004: 219-221.
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XIAO S H, YAN C S, ZHANG H P, SUN G Z. Study on Wheat Spike Germination. Beijing: China Agricultural Science and Technology Press, 2004: 219-221. (in Chinese)
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[26] 国家质量技术监督局. 小麦和小麦粉面筋含量第2部分:仪器法测定湿面筋: GB/T 5506.2- 2008. 北京: 中国标准出版社, 2008.
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[27] 国家质量技术监督局. 小麦、黑麦及其面粉、杜伦麦及其粗粒粉降落数值的测定: GB/T 10361-2008. 北京: 中国标准出版社, 2008.
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[28] 国家质量技术监督局. 小麦、黑麦及其粉类和淀粉糊化特性测定快速粘度仪法: GB/T 24853-2010. 北京: 中国标准出版社, 2010.
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[30] 刘一灵, 李振华, 刘仁祥 . 植物激素ABA调控种子发育与萌发的研究进展
种子, 2014,33(10):47-50.
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LIU Y N, LI Z H, LIU R X . The research of ABA regulation in seed development and germination
Seed, 2014,33(10):47-50. (in Chinese)
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[31] 杨剑平 . 拟南芥种子脱水过程中ABA作用的研究综述
湖北农学院学报, 1995,15(2):133-136.
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YANG J P . Review on the role of ABA in Arabidopsis seed dehydration
Journal of Hubei Agricultural College, 1995,15(2):133-136. (in Chinese)
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[32] 孙果忠, 张秀英, 闫长生, 肖世和 . 不同穗发芽抗性的小麦胚对ABA敏感性及抗性机制研究
麦类作物学报, 2005,25(2):27-32.
DOI:10.7606/j.issn.1009-1041.2005.02.053URL [本文引用: 2]
以20个小麦品种为材料,测定了不同穗发芽抗性的小麦品种处于种子发育过程中的胚对ABA敏感性和后熟过程中种子的萌发能力,以揭示小麦对穗发芽的抗性机制。结果表明,在种子发育过程中,不同抗性品种的胚萌发能力和胚对ABA敏感性存在差异。胚萌发能力随胚龄的增加,呈"低～高～低"的变化趋势。扬花后25d到35d的胚萌发势的降幅值在抗穗发芽基因型中所占的比重明显较易穗发芽基因型大。易穗发芽基因型对ABA的敏感性随胚萌发能力的提高而下降,而抗穗发芽的基因型对ABA的敏感性随胚萌发能力的提高则有升有降。因此,胚休眠主要在发育后期获得。红皮易穗发芽品种的成熟胚因脱离母体而提高的萌发势明显较白皮易穗发芽品种的高。品种的成熟胚离体萌发能力越高,则种子休眠期越短。
SUN G Z, ZHANG X Y, YAN C S, XIAO S H . Embryonic ABA sensitivity and resistant mechanism of sprouting resistance and susceptible cultivars
Journal of Triticeae Crop, 2005,25(2):27-32. (in Chinese)
DOI:10.7606/j.issn.1009-1041.2005.02.053URL [本文引用: 2]
以20个小麦品种为材料,测定了不同穗发芽抗性的小麦品种处于种子发育过程中的胚对ABA敏感性和后熟过程中种子的萌发能力,以揭示小麦对穗发芽的抗性机制。结果表明,在种子发育过程中,不同抗性品种的胚萌发能力和胚对ABA敏感性存在差异。胚萌发能力随胚龄的增加,呈"低～高～低"的变化趋势。扬花后25d到35d的胚萌发势的降幅值在抗穗发芽基因型中所占的比重明显较易穗发芽基因型大。易穗发芽基因型对ABA的敏感性随胚萌发能力的提高而下降,而抗穗发芽的基因型对ABA的敏感性随胚萌发能力的提高则有升有降。因此,胚休眠主要在发育后期获得。红皮易穗发芽品种的成熟胚因脱离母体而提高的萌发势明显较白皮易穗发芽品种的高。品种的成熟胚离体萌发能力越高,则种子休眠期越短。

[33] 周述波, 贺立红, 林伟, 戴高兴 . 外源GA3、ABA对杂交水稻亲本穗上发芽的生理影响
湖北农业科学, 2011,50(21):19-22.
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ZHOU S B, HE L H, LIN W, DAI G X . Effects of exogenous GA3 and ABA on physiological change of preharvest sprouting in hybrid rice
Hubei Agricultural Sciences, 2011,50(21):19-22. (in Chinese)
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[34] 吴颖, 胡汉桥, 王罡, 张艳贞, 季静 . 春小麦α-淀粉酶活性及其与穗发芽抗性的关系
吉林农业大学学报, 2002,24(4):22-25.
URL [本文引用: 1]
The pre-harvest sprouting resistance of 57 spring wheat varieties (lines) were determined in 35 d periods after the anthesis, and the α-amylase activity was also determined by the improved method of gel diffusion. The results showed that differences between pre-harvest sprouting resistance and α-amylase activity of these varieties were significant. Most varieties of Heilongjiang province had only middle resistance to pre-harvest sprouting. The pre-harvest sprouting resistance of red color coat materials was higher than that of white materials, and there were highly resistant varieties among these white color varieties. Pre-harvest sprouting rates were positively correlated with α-amylase concentrations. α-amylase concentrations in 35 d after anthesis could be a criterion of pre-harvest sprouting resistance.
WU Y, HU H Q, WANG G, ZHANG Y Z, JI J . Relationship between α-amylase activity and resistance of pre-harvest sprouting in spring wheat
Journal of Jilin Agricultural University, 2002,24(4):22-25. (in Chinese)
URL [本文引用: 1]
The pre-harvest sprouting resistance of 57 spring wheat varieties (lines) were determined in 35 d periods after the anthesis, and the α-amylase activity was also determined by the improved method of gel diffusion. The results showed that differences between pre-harvest sprouting resistance and α-amylase activity of these varieties were significant. Most varieties of Heilongjiang province had only middle resistance to pre-harvest sprouting. The pre-harvest sprouting resistance of red color coat materials was higher than that of white materials, and there were highly resistant varieties among these white color varieties. Pre-harvest sprouting rates were positively correlated with α-amylase concentrations. α-amylase concentrations in 35 d after anthesis could be a criterion of pre-harvest sprouting resistance.

[35] 张笑, 赵纯钦, 黄静, 徐登安, 张赤红, 陈静 . 外源脱落酸、水杨酸对小麦种子萌发及生理特性的影响
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Chinese Journal of Applied & Environmental Biology, 2014,20(1):139-143. (in Chinese)
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[36] 张正斌, 谈成, 徐明华, 牟会荣 . 外源脱落酸和冠菌素对小麦籽粒萌芽的影响
安徽农业科学, 2013,41(13):5649-5652.
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ZHANG Z B, TAN C, XU M H, MOU H R . Effects of abscisic acid and coronatine on sprouting of winter wheat grain
Journal of Anhui Agricultural Sciences, 2013,41(13):5649-5652. (in Chinese)
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[37] 任江萍, 陈焕丽, 王振云, 尹钧, 李永春, 牛洪斌, 宋晓 . 小麦穗发芽与籽粒内可溶性糖和α-淀粉酶活性的品种差异
西北农业科学, 2007,16(1):22-25.
[本文引用: 1]

REN J P, CHEN H L, WANG Z Y, YIN J, LI Y C, NIU H B, SONG X . Difference of pre-harvest sprouting and soluble sugar content and α-amylase activity in grains of some wheat varieties during grain developing
Acta Agriculturae Boreali-Occidentalis Sinica, 2007,16(1):22-25. (in Chinese)
[本文引用: 1]

[38] STAROSKE N, UDO C, KUMLEHN J, HENSEL G, RADCHUK R, ERBAN A, KOPKA J, WESCHKE W, WEBER H . Increasing abscisic acid levels by immunomodulation in barley grains induces precocious maturation without changing grain composition
Journal of Experimental Botany, 2016,67(9):2675-2687.
DOI:10.1093/jxb/erw102URLPMID:26951372 [本文引用: 1]
Abscisic acid (ABA) accumulates in seeds during the transition to the seed filling phase. ABA triggers seed maturation, storage activity, and stress signalling and tolerance. Immunomodulation was used to alter the ABA status in barley grains, with the resulting transgenic caryopses responding to the anti-ABA antibody gene expression with increased accumulation of ABA. Calculation of free versus antibody-bound ABA reveals large excess of free ABA, increasing signficantly in caryopses from 10 days after fertilization. Metabolite and transcript profiling in anti-ABA grains expose triggered and enhanced ABA-functions such as transcriptional up-regulation of sucrose-to-starch metabolism, storage protein synthesis and ABA-related signal transduction. Thus, enhanced ABA during transition phases induces precocious maturation but negatively interferes with growth and development. Anti-ABA grains display broad constitutive gene induction related to biotic and abiotic stresses. Most of these genes are ABA- and/or stress-inducible, including alcohol and aldehyde dehydrogenases, peroxidases, chaperones, glutathione-S-transferase, drought- and salt-inducible proteins. Conclusively, ABA immunomodulation results in precocious ABA accumulation that generates an integrated response of stress and maturation. Repression of ABA signalling, occurring in anti-ABA grains, potentially antagonizes effects caused by overshooting production. Finally, mature grain weight and composition are unchanged in anti-ABA plants, although germination is somewhat delayed. This indicates that anti-ABA caryopses induce specific mechanisms to desensitize ABA signalling efficiently, which finally yields mature grains with nearly unchanged dry weight and composition. Such compensation implicates the enormous physiological and metabolic flexibilities of barley grains to adjust effects of unnaturally high ABA amounts in order to ensure and maintain proper grain development.

[39] 赵虎成, 王伯初, 刘堰, 蔡绍皙 . 逆境胁迫下植物基因的表达与调控
重庆大学学报(自然科学版), 2000,23(5):146-148.
[本文引用: 1]

ZHAO H C, WANG B C, LIU Y, CAI S X . Expression and regulation of the plant gene under enviro mend stress
Journal of Chongqing University(Natural Science Edition), 2000,23(5):146-148. (in Chinese)
[本文引用: 1]

[40] 刘锐, 魏益民, 邢亚楠, 张波, 张影全 . 小麦淀粉与面条质量关系的研究进展
麦类作物学报, 2013,33(5):1058-1063.
DOI:10.7606/j.issn.1009-1041.2013.05.037URL [本文引用: 1]
Starch quality has great influence on white salted noodles quality, especially on the sensory characteristics of cooked noodles. The contents of amylose and amylopectin result in differences in swelling power and pasting properties of starch, and thus significantly affect noodle quality. Wheat flours with lower amylose content have better pasting and swelling properties, and the noodles prepared from this flour have more water absorption, lower cooking loss and higher sensory scores. Optimum amylose content of wheat flour with good noodle making quality is around 22%. Peak viscosity, breakdown, and peak time are important pasting parameters affecting noodles quality. Wheat flours with high peak viscosity, breakdown and peak time appear to be suitable for noodle making. A soft, smooth and springy texture is found in noodles produced from wheat flour with high starch swelling power or swelling volume. Generally, low amylose content, high past peak viscosity, peak time and breakdown, and high starch swelling power or swelling volume have been reported as desirable for good noodles quality. Amylose content, peak viscosity and swelling power are the key characteristics in the evaluation of suitability of wheat flour for making noodles.
LIU R, WEI Y M, XING Y N, ZHANG B, ZHANG Y Q . Review on the relationship between starch and noodle quality in wheat
Journal of Triticeae Crops, 2013,33(5):1058-1063. (in Chinese)
DOI:10.7606/j.issn.1009-1041.2013.05.037URL [本文引用: 1]
Starch quality has great influence on white salted noodles quality, especially on the sensory characteristics of cooked noodles. The contents of amylose and amylopectin result in differences in swelling power and pasting properties of starch, and thus significantly affect noodle quality. Wheat flours with lower amylose content have better pasting and swelling properties, and the noodles prepared from this flour have more water absorption, lower cooking loss and higher sensory scores. Optimum amylose content of wheat flour with good noodle making quality is around 22%. Peak viscosity, breakdown, and peak time are important pasting parameters affecting noodles quality. Wheat flours with high peak viscosity, breakdown and peak time appear to be suitable for noodle making. A soft, smooth and springy texture is found in noodles produced from wheat flour with high starch swelling power or swelling volume. Generally, low amylose content, high past peak viscosity, peak time and breakdown, and high starch swelling power or swelling volume have been reported as desirable for good noodles quality. Amylose content, peak viscosity and swelling power are the key characteristics in the evaluation of suitability of wheat flour for making noodles.

[41] SEIB P A . Reduced-amylose wheats and Asian noodles
Cereal Foods World, 2000,45(11):504-512.
[本文引用: 1]

[42] 文廷刚, 于洪喜, 王伟中, 杨文飞, 钱新民 . 不同生长期喷施植物生长调节剂对小麦籽粒品质的影响
金陵科技学院学报, 2017,33(4):53-57.
[本文引用: 1]

WEN Y G, YU H X, WANG W Z, YANG W F, QIAN X M . Effects of PGRs treatment applied at different growth periods on grain quality in wheat
Journal of Jinling Institute of Technology, 2017,33(4):53-57. (in Chinese)
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[43] 王枟刘, 闫素辉, 王其飞, 陶胜, 刘颂 . 施氮水平对稻茬晚播小麦籽粒淀粉含量与糊化特性的影响
安徽科技学院学报, 2015(3):6-9.
[本文引用: 1]

WANG Y L, YAN S H, WANG Q F, TAO S, LIU S . Effects of nitrogen rate on starch content and pasting properties in grain of late sowing wheat in rice-wheat rotation
Journal of Anhui Science and Technology University, 2015(3):6-9. (in Chinese)
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[44] 马丽荣 . 马铃薯淀粉代谢相关基因表达与淀粉品质研究
[D]. 银川: 宁夏大学, 2015.
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MA L R . Study on gene expression involving in starch metabolism and starch quality in potato
[D]. Yinchuan: Ningxia University, 2015. ( in Chinese)
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[45] YANG J C, ZHANG J H, WANG Z Q, ZHU Q S . Activities of key enzymes in sucrose-to-starch conversion in wheat grains subjected to water deficit during grain filling
Plant Physiology, 2004,135(3):1621-1629.
DOI:10.1104/pp.104.041038URLPMID:15235118 [本文引用: 1]
This study tested the hypothesis that a controlled water deficit during grain filling of wheat (Triticum aestivum) could accelerate grain-filling rate through regulating the key enzymes involved in Suc-to-starch pathway in the grains. Two high lodging-resistant wheat cultivars were field grown. Well-watered and water-deficit (WD) treatments were imposed from 9 DPA until maturity. The WD promoted the reallocation of prefixed 14C from the stems to grains, shortened the grain-filling period, and increased grain-filling rate or starch accumulation rate (SAR) in the grains. Activities of Suc synthase (SuSase), soluble starch synthase (SSS), and starch branching enzyme (SBE) in the grains were substantially enhanced by WD and positively correlated with the SAR. ADP Glc pyrophosphorylase activity was also enhanced in WD grains initially and correlated with SAR with a smaller coefficient. Activities of granule-bound starch synthase and soluble and insoluble acid invertase in the grains were less affected by WD. Abscisic acid (ABA) content in the grains was remarkably enhanced by WD and very significantly correlated with activities of SuSase, SSS, and SBE. Application of ABA on well-watered plants showed similar results as those by WD. Spraying with fluridone, an ABA synthesis inhibitor, had the opposite effect. The results suggest that increased grain-filling rate is mainly attributed to the enhanced sink activity by regulating key enzymes involved in Suc-to-starch conversion, especially SuSase, SSS, and SBE, in wheat grains when subjected to a mild water deficit during grain filling, and ABA plays a vital role in the regulation of this process.

[46] YANG J C, ZHANG J H, WANG Z Q, ZHU Q S . Hormonal changes in the grains of rice subjected to water stress during grain filling
Plant Physiology, 2001,127(1):315-323.
DOI:10.1104/pp.127.1.315URLPMID:11553759 [本文引用: 1]
Lodging-resistant rice (Oryza sativa) cultivars usually show slow grain filling when nitrogen is applied in large amounts. This study investigated the possibility that a hormonal change may mediate the effect of water deficit that enhances whole plant senescence and speeds up grain filling. Two rice cultivars showing high lodging resistance and slow grain filling were field grown and applied with either normal or high amount nitrogen (HN) at heading. Well-watered and water-stressed (WS) treatments were imposed 9 days post anthesis to maturity. Results showed that WS increased partitioning of fixed (14)CO(2) into grains, accelerated the grain filling rate but shortened the grain filling period, whereas the HN did the opposite way. Cytokinin (zeatin + zeatin riboside) and indole-3-acetic acid contents in the grains transiently increased at early filling stage and WS treatments hastened their declines at the late grain filling stage. Gibberellins (GAs; GA(1) + GA(4)) in the grains were also high at early grain filling but HN enhanced, whereas WS substantially reduced, its accumulation. Opposite to GAs, abscisic acid (ABA) in the grains was low at early grain filling but WS remarkably enhanced its accumulation. The peak values of ABA were significantly correlated with the maximum grain filling rates (r = 0.92**, P < 0.01) and the partitioning of fixed (14)C into grains (r = 0.95**, P < 0.01). Exogenously applied ABA on pot-grown HN rice showed similar results as those by WS. Results suggest that an altered hormonal balance in rice grains by water stress during grain filling, especially a decrease in GAs and an increase in ABA, enhances the remobilization of prestored carbon to the grains and accelerates the grain filling rate.

[47] CHEN K, LI G J, BRESSAN R, SONG C P . Abscisic acid dynamics, signaling, and functions in plants
Journal of Integrative Plant Biology, 2020,62(1):25-54.
DOI:10.1111/jipb.12899URLPMID:31850654 [本文引用: 1]
Abscisic acid (ABA) is an important phytohormone regulating plant growth, development, and stress responses. It has an essential role in multiple physiological processes of plants, such as stomatal closure, cuticular wax accumulation, leaf senescence, bud dormancy, seed germination, osmotic regulation, and growth inhibition among many others. Abscisic acid controls downstream responses to abiotic and biotic environmental changes through both transcriptional and posttranscriptional mechanisms. During the past 20 years, ABA biosynthesis and many of its signaling pathways have been well characterized. Here we review the dynamics of ABA metabolic pools and signaling that affects many of its physiological functions.

[48] PENG D L, CAI T, YIN Y P, YANG W B, NI Y L, YANG D Q, WANG Z L . Exogenous application of abscisic acid or gibberellin acid has different effects on starch granule size distribution in grains of wheat
Journal of Integrative Agriculture, 2013,12(9):1551-1559.
DOI:10.1016/S2095-3119(13)60557-2URL [本文引用: 1]
Granule size distribution of wheat starch is an important characteristic that can affect its chemical composition and the functionality of wheat products. Two high-yield winter wheat cultivars were used to evaluate the effects of the application of exogenous ABA or GA during the reproductive phase of the initial grain filling on starch granule size distribution and starch components in grains at maturity. The results indicated that a bimodal curve was found in the volume and surface area distribution of grain starch granules, and a unimodal curve was observed for the number distribution under all treatments. The exogenous ABA resulted in a significant increase in the proportions (both by volume and by surface area) of B-type (<9.9 mu m in diameter) starch granules, with a reduction in those of A-type (>9.9 mu m) starch granules, while, the exogenous GA(3) led to converse effects on size distribution of those starch granules. The exogenous ABA also increased starch, amylose and amylopectin contents at maturity but significantly reduced the ratio of amylose to amylopectin. Application of GA(3) significantly reduced starch content, amylopectin content but increased the ratio of amylose to amylopectin. The ratio of amylose to amylopectin showed a significant and negative relationship with the volume proportion of granules <9.9 gm, but was positively related to the volume proportion of granules 22.8-42.8 mu m.