花后DA-6处理调控小麦种子活力的机理

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许晨^,, 王文静, 曹珊, 李如雪, 张贝贝, 孙爱清^,, 张春庆^,山东农业大学农学院/作物生物学国家重点实验室/山东省作物生物学重点实验室,山东泰安 271018

Mechanism of DA-6 Treatment Regulating Wheat Seed Vigor After Anthesis

XU Chen^,, WANG WenJing, CAO Shan, LI RuXue, ZHANG BeiBei, SUN AiQing^,, ZHANG ChunQing^,College of Agriculture, Shandong Agricultural University/State Key Laboratory of Crop Biology/Shandong Key Laboratory of Crop Biology, Tai’an 271018, Shandong

通讯作者: 张春庆,E-mail：cqzhang@sdau.edu.cn。孙爱清,E-mail：saqsshh@sdau.edu.cn

责任编辑: 李莉
收稿日期:2020-08-24接受日期:2020-10-19网络出版日期:2021-05-01

基金资助:

国家重点研发计划七大农作物育种专项课题.2018YFD0100904

Received:2020-08-24Accepted:2020-10-19Online:2021-05-01
作者简介 About authors
许晨,E-mail：xuchen0905@foxmail.com。

摘要
【背景】高活力的种子萌发迅速整齐,抗逆性强,具有很大的生产优势。己酸二乙氨基乙醇酯（DA-6）作为叔胺类植物生长调节剂,可以提高植物的光合速率,调节碳氮代谢,提高作物品质和产量。【目的】分析花后喷施DA-6对小麦籽粒物质积累、物质代谢和抗逆关键基因表达、种子千粒重、化学成分与种子活力的影响,明确DA-6调控小麦种子活力的机理,为DA-6在小麦高活力种子生产调控中的应用提供理论依据和技术支持。【方法】以小麦品种良星77和山农23号为材料,于花后2和6 d进行DA-6田间喷施。试验于2017—2018年、2018—2019年在山东农业大学马庄试验基地（117°E;36°N）进行,设置2个处理浓度（0和6 g·L^-1 DA-6）和2个处理时期（花后2和6 d）,设置3个区组,每个小区面积6 m×3 m。于籽粒灌浆前期和中期（花后12和22 d）取籽粒、穗下节和旗叶鲜样,用于物质代谢关键酶基因表达的检测。于籽粒发育的不同天数取样（花后25、30、35和40 d）,自然晒干后脱粒测定千粒重和蛋白质含量、淀粉含量等;对于成熟期收获的小麦穗,取小麦整穗、不同穗位（上部、中部和下部）、不同粒位（下位粒和上位粒）的种子测定发芽活力指标。【结果】花后6 d喷施6 g·L^-1DA-6可大幅度提高千粒重及种子蛋白质含量,使2个小麦品种的穗上部和上位粒种子的单株干重显著增加,显著提高种子的活力指数。花后6 d喷施DA-6有效降低了不同穗粒位之间的种子活力差异,使小麦不同穗位、粒位种子的活力得到整体提高,尤其以穗上中部籽粒及上位粒的种子活力提高更为明显。而花后2 d喷施DA-6则使种子活力显著降低,千粒重及种子蛋白质含量也有降低。良星77和山农23号的千粒重均表现为随着籽粒灌浆期的延长,千粒重先升高后降低;但相对蛋白及绝对蛋白含量则表现为随着时间的延长而渐升。同时,两品种均表现为花后6 d喷施6 g·L^-1的DA-6时千粒重及蛋白质含量最高。此处理下,良星77花后40 d收获时,千粒重与对照相比无显著差异,但相对蛋白质含量、绝对蛋白质含量分别较对照显著提高;山农23号花后6 d喷施6 g·L^-1 DA-6后千粒重、绝对蛋白含量均显著提高。两品种花后2 d喷施DA-6对千粒重、相对蛋白质含量、绝对蛋白含量及淀粉含量无显著影响。实时荧光定量PCR表明,花后6 d喷施DA-6可诱导叶片蛋白磷酸酶和蔗糖磷酸合成酶基因表达水平显著上调,茎秆中Ⅰ类hsp和hsp70表达量显著提高,籽粒中蛋白质二硫键异构酶和hsp82的表达也显著上调。花后6 d喷施DA-6处理的小麦种子萌发期α-淀粉酶活性检测表明,山农23号的对照和处理组均表现为其α-淀粉酶活性随着萌发时间的延长逐渐增强,喷施DA-6可显著提高2个品种的种子萌发72 h的α-淀粉酶活性。【结论】花后6 d喷施DA-6促进小麦中蛋白和蔗糖代谢关键酶基因及热激蛋白基因的表达,有利于促进种子贮藏物质合成,使种子蛋白质含量升高,最终导致千粒重提高;同时还显著增加2个小麦品种穗上中部及上位粒种子萌发过程中的单株干重,提高种子的活力指数,降低不同穗粒位之间种子活力的差异,提高种子萌发过程中的α-淀粉酶活性和物质转化能力,促进幼苗干物质积累,最终提高种子活力。小麦种子花后6 d喷施6 g·L^-1 DA-6是提高小麦种子活力的有效途径。
关键词： 小麦;己酸二乙氨基乙醇酯;种子活力;实时荧光定量PCR

Abstract

【Background】High-vigor seeds germinate quickly and efficiently, and they are highly stress resistant and advantageous during crop production. Diethyl aminoethyl hexanoate (DA-6), as a tertiary amine plant growth regulator, increases plant photosynthetic rates, regulates carbon and nitrogen metabolism and improves crop quality and yield. However, there are limited studies on its effects on seed vigor. 【Objective】This study analyzed the effects of spraying DA-6 post-flowering on the compounds accumulated in grains, expression levels of key genes associated with metabolism and stress-resistance, 1000-grain weights, chemical compositions and seed vigor to determine DA-6’s mechanism for regulating wheat seed vigor, which has important theoretical and practical significance in the production of high-vigor wheat seeds. 【Method】Wheat varieties, Liangxing 77 and Shannong 23, grown in fields of the Mazhuang Experimental Base of Shandong Agricultural University in 2017-2018 and 2018-2019 (117°E, 36°N) were sprayed with DA-6. Two treatment concentrations (0 and 6 g·L^-1 DA-6), two treatment periods (2 and 6 days after anthesis) and three zone groups, each having an area of 6 m × 3 m, were used. Fresh samples of grains, stems below spikes and flag leaves were collected at the early and middle stages of grain filling (12 and 22 days after anthesis, respectively), and the expression levels of key metabolism-related enzymes were analyzed. Samples were taken on different days of grain development (25, 30, 35 and 40 days after anthesis), and the 1000-grain weights, protein contents and starch contents were determined after natural drying and threshing. For wheat spikes harvested at the maturity stage, the germination and vigor indices of the whole ear, different spike positions (upper, middle and lower) and different grain positions (lower and upper grain) were measured. 【Result】Spraying 6 g·L^-1 DA-6 at 6 days after anthesis significantly increased the 1000-grain weights, seed protein contents, and plant dry weights of the upper spikes and upper grains in both wheat varieties, and it significantly improved the seed vigor indices. DA-6 spraying at 6 days after anthesis effectively reduced the seed vigor difference between different spike and grain positions, and it improved the seed vigor of different spike and grain positions in wheat as a whole, especially those of upper and middle spikes, as well as the upper grain. However, DA-6 spraying at 2 days after anthesis significantly reduced seed vigor, 1000-grain weights and seed protein contents. The 1000-grain weights of Liangxing 77 and Shannong 23 increased first and then decreased as the grain-filling period was extended. However, relative and absolute protein contents increased with time. Additionally, both varieties showed the highest 1000-grain weights and protein contents after spraying 6 g·L^-1 DA-6 at 6 days after anthesis. While the 1000-grain weight of Liangxing 77 was not significantly different from that of the control group, the relative and absolute protein contents were significantly greater. The 1000-grain weight and absolute protein content of Shannong 23 significantly increased after spraying 6 g·L^-1 DA-6 at 6 days after anthesis. DA-6 spraying at 2 days after anthesis had no significant effects on 1000-grain weights, relative and absolute protein contents or starch contents. Real fluorescence quantitative PCR showed that DA-6 spraying at 6 days after anthesis significantly increased the expression levels of protein phosphatase and sucrose phosphate synthase in flag leaves. The expression levels of class I heat shock protein (hsp) and hsp70 in the stem, like those of protein disulfide isomerase and hsp82 in grains, increased significantly. The α-amylase activity during seed germination was measured in samples sprayed with DA-6 at 6 days after anthesis. The activity gradually increased along with germination time in both control and treated Shannong 23. DA-6 spraying at 6 days after anthesis significantly increased the α-amylase activity levels in seeds of both varieties after germinating for 72 h. 【Conclusion】Spraying DA-6 at 6 days after anthesis promoted the expression levels of proteins, key enzyme-encoding genes of sucrose metabolism and hsps. It also facilitated the synthesis of seed storage substances, increased the protein contents of seeds and significantly increased the 1000-grain weights. The dry weight of individual plants during germination and seed vigor of upper, middle spikes and the upper grain of harvested seeds in two varieties significantly improved after spraying DA-6. The seed vigor difference between different spike positions decreased, and the α-amylase activity increased during seed germination. Thus, spraying DA-6 after anthesis promoted the accumulation of seedling dry matter and improved material transformation capacities during seed germination. Additionally, seed vigor improved. Our studies suggest that spraying 6 g·L^-1 DA-6 on wheat at 6 days after anthesis is an effective way to improve seed vigor.

Keywords：wheat;DA-6;seed vigor;real-time PCR

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本文引用格式
许晨, 王文静, 曹珊, 李如雪, 张贝贝, 孙爱清, 张春庆. 花后DA-6处理调控小麦种子活力的机理[J]. 中国农业科学, 2021, 54(9): 1821-1834 doi:10.3864/j.issn.0578-1752.2021.09.001
XU Chen, WANG WenJing, CAO Shan, LI RuXue, ZHANG BeiBei, SUN AiQing, ZHANG ChunQing. Mechanism of DA-6 Treatment Regulating Wheat Seed Vigor After Anthesis[J]. Scientia Acricultura Sinica, 2021, 54(9): 1821-1834 doi:10.3864/j.issn.0578-1752.2021.09.001

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0 引言

【研究意义】种子是农业生产的重要基础,种子的品种品质和播种品质的高低则是农业健康稳定发展的关键,而种子活力更是重中之重。高活力种子不仅快速突破种皮,使出苗整齐,而且还具有较强的抗逆性,产生明显的生长优势和生产潜力^[1,2,3,4]。己酸二乙氨基乙醇酯（N,N-diethyl aminoethyl hexanoate,DA-6）是一种具有较高生物活性、人工合成的植物生长调节剂,俗称胺鲜酯^[5,6],目前,DA-6已被广泛应用于农作物、果蔬、花卉等领域^{[7,8,9,10,11]},具有增产、抗逆、抗病、改善品质等功效。因此,研究不同浓度DA-6对种子活力及产量的调控机理,对利用植物生长调节剂处理提高小麦种子活力及产量具有重要的理论意义和实践价值。【前人研究进展】关于DA-6对种子活力及产量的影响,前人用不同方法做了大量研究。XIE等^[12]研究发现外源喷施DA-6可以提高玉米根系导水率、促进幼苗的生长,增加植株叶面积,增加根和叶干重。而在其他作物上如番茄^[13]、辣椒^[14]、艾纳香^[15]、生菜^[16]等喷施DA-6可促进生长,增加干重。聂乐兴等^[17,18]研究发现拔节期喷施DA-6增产效果明显,处理后玉米的叶面积指数、光合速率、RuBP羧化酶和PEP羧化酶活性均显著上升。ZHOU等^[19]研究发现200 μmol·L^-1的DA-6可通过提高脂肪酸和甘油向糖类的转化而促进老化大豆种子幼苗的形态建成,提高种子活力。研究还发现DA-6浸种可提高番茄^[20]、水稻^[21,22]、玉米^[23,24,25]等作物的发芽率及活力指数。WEN等^[26]对小麦和玉米种子活力的研究表明种子内的蛋白质含量与种子活力显著相关。多项研究表明在小麦、大麦、燕麦等作物中种子活力与种子大小和蛋白质含量显著相关^{[27,28,29,30]}。【本研究切入点】前人研究主要采用外源喷施DA-6提高作物产量、改善作物品质,采用DA-6溶液浸种的方式提高种子活力,且多集中于玉米、水稻等作物,而外源喷施DA-6提高小麦种子活力的研究较少。【拟解决的关键问题】本研究通过测定DA-6喷施处理后小麦生长进程中粒重、化学成分、籽粒物质代谢关键酶基因表达的变化,收获后不同穗位、粒位种子发芽活力的差异,以期明确DA-6调控小麦种子活力和产量形成的机理,为DA-6在小麦高活力种子生产调控中的应用提供理论依据和技术支持。

1 材料与方法

1.1 材料

小麦品种良星77购自山东良星种业有限公司,半冬性,多穗型;山农23号购自济南鑫瑞种业科技有限公司,半冬性,大穗型。己酸二乙氨基乙醇酯粉剂（DA-6）,有效含量为98.5%,由常州金坛茂盛精细化工厂生产。DA-6溶液配制时添加0.1%有机硅助剂（液体,河北农迅生物科技有限公司生产）。

1.2 试验设计

试验于2017—2018年、2018—2019年在山东农业大学马庄试验基地（117°E;36°N）进行,常规栽培管理。田间试验采用随机区组设计,每个处理3个区组,花后喷施DA-6。

2017—2018年,选择1个小麦品种（良星77）,设置2个处理浓度（0和6 g·L^-1 DA-6）、2个处理时期（花后2 d、花后6 d）和3个区组,小区面积为6 m×3 m。将良星77的盛花期记为开花0 d,分别于花后2 d（2018年4月26日）、花后6 d（4月30日）进行DA-6喷施处理,喷施量为750 L·hm^-2。于籽粒发育的不同天数取样（开花后25、30、35和40 d）,每次取样1 m²,自然晒干后测定粒重和蛋白质含量、淀粉含量等;对成熟期收获的小麦穗,取小麦整穗、不同穗位（上部、中部和下部）、不同粒位（下位粒和上位粒）的种子测定发芽活力指标。

2018—2019年,选择2个小麦品种（良星77和山农23号）,设置2个处理浓度（0和6 g·L^-1 DA-6）、2个处理时期（花后2和6 d）和3个区组,小区面积为6 m×3 m。两品种各自盛花期记为开花0 d,分别于花后2 d（良星77为2019年5月2日,山农23号为5月3日）、花后6 d（良星77为5月6日,山农23号为5月7日）进行DA-6喷施处理,喷施量为1 800 L·hm^-2。于籽粒发育的不同天数取样（花后25、30、35和40 d）,每次取样1 m²,自然晒干后脱粒测定粒重和蛋白质含量、淀粉含量等;对成熟期收获的小麦穗,取小麦整穗、不同穗位（上部、中部和下部）、不同粒位（下位粒和上位粒）的种子,测定发芽活力指标。于籽粒灌浆前期和中期（花后12和22 d）取样,分别取籽粒、穗下节和旗叶的鲜样,迅速液氮冷冻,-80℃保存,用于物质代谢关键酶基因表达的检测。

1.3 千粒重测定

参照国家标准法（GB 5519-2008）,测定小麦千粒重,据种子实际含水量换算至标准水分（13%）条件下的千粒重,3次重复,取平均值。

标准水分（13%）千粒重（g）=实测千粒重（g）×（1-实测含水量,%）/（1-13%）。

1.4 籽粒化学成分

采用Perten DA7200近红外谷物品质分析仪测定籽粒的含水量、蛋白质、淀粉百分数含量,换算为规定水分（13%）条件下的化学成分含量,3次重复,取平均值。

相对蛋白质含量（relative content of protein,RCP,%）=实测蛋白质%×（1-13%）/（1-实测含水量,%）;

绝对蛋白质含量（absolute content of protein,ACP,mg/seed）=实测千粒重（g）×实测蛋白质（%）/100;

相对淀粉含量（relative content of starch,RCS,%）=实测淀粉（%）×（1-13%）/（1-实测含水量,%）;

绝对淀粉含量（absolute content of starch,ACS,mg/seed）=实测千粒重（g）×实测淀粉（%）/100。

1.5 种子发芽活力指标的测定

参照国家标准农作物种子检验规程（GB/T 3543.4-1995）,进行标准发芽试验,测定种子发芽势（germination energy,GE）、发芽率（germination percentage,GP）、发芽指数（germination index,GI）和活力指数（vigor index,VI）。其中,VI=GI×S,S为一定时期内正常幼苗单株干重。

1.6 α-淀粉酶活性的测定

采用3,5-二硝基水杨酸（3,5-Dinitrosalicylic acid,DNS）法测定α-淀粉酶活性^[31,32],略作改进。

1.7 RNA提取及qRT-PCR

采用天根公司DP441试剂盒（天根,中国北京）提取总RNA,用TaKaRa试剂盒（PrimeScript RT reagent Kit With gDNA Eraser）合成cDNA,ABI Stepone plus荧光定量PCR仪进行荧光定量PCR,参考WEN等^[33]方法设计引物（表1）。

Table 1
表1
表1qRT-PCR引物
Table 1Primers used for qRT-PCR

基因Gene	引物名称Primer name	序列Sequence (5′-3′)
肌动蛋白基因（内参）Actin gene (internal reference)	Actin-F	CGAAGCGACATACAATTCCATC
肌动蛋白基因（内参）Actin gene (internal reference)	Actin-R	GAACCTCCACTGAGAACAACAT
蛋白磷酸酶基因Protein phosphatase gene	TraesCS3B02G277900-F	CTGGCAGGTGGTTGGAGGAAT
蛋白磷酸酶基因Protein phosphatase gene	TraesCS3B02G277900-R	GCAATAGGCTCAGGACGGATCT
蔗糖磷酸合成酶基因Sucrose phosphate synthase gene	TraesCS3A02G015500-F	GTGCCAATGGTCTTCACAGGTC
蔗糖磷酸合成酶基因Sucrose phosphate synthase gene	TraesCS3A02G015500-R	TCAGCCTCAATCCGTCGCATTA
蛋白质二硫键异构酶基因Protein disulfide isomerase gene	TraesCS4A02G214200-F	CCCTGACAACCATCCTTACCTCT
蛋白质二硫键异构酶基因Protein disulfide isomerase gene	TraesCS4A02G214200-R	CCTTGCCACTGAACTCCTCTACA
hsp82	TraesCS2A02G033700-F	GCTGCTCTCGCTCATCATCAAC
hsp82	TraesCS2A02G033700-R	TGCTCTTGTCTGTCAGGCTCTC
Ⅰ类hsp ClassⅠ hsp	TraesCS4B02G211700-F	ACAAGGAGCAGGAGGAGAAGAC
Ⅰ类hsp ClassⅠ hsp	TraesCS4B02G211700-R	TGACGGTGAGCACGCTGTT
hsp70	TraesCS1D02G284000-F	CGGCATCCTGAACGTGTCTG
hsp70	TraesCS1D02G284000-R	TCTTCTTGTGCTCCTCGTCCTC

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1.8 数据分析

采用Microsoft Excel 2013计算试验数据平均值并作图,用SPSS 21.0进行数据统计分析,LSD法比较处理之间的差异显著程度。

2 结果

2.1 喷施DA-6对不同穗位、粒位种子发芽活力的影响

标准发芽试验结果表明,2018年花后6 d喷施DA-6显著提高了良星77穗上部种子的单株干重及活力指数,分别为3.92%和5.22%。良星77花后2 d喷施DA-6,显著降低了穗上部和穗中部种子的单株干重及活力指数,其中,穗上部种子的单株干重与活力指数与对照相比分别显著降低4.49%和2.87%,穗中部种子的单株干重与活力指数与对照相比分别显著降低7.11%和8.37%。对不同粒位的小麦来说,花后6 d喷施DA-6使上位粒的单株干重及活力指数显著提高,分别提高8.54%和10.61%;对下位粒的单株干重无显著影响,但可使其活力指数显著提高3.37%。花后2 d喷施DA-6使下位粒的种子活力显著降低,与对照相比,单株干重和活力指数分别显著降低3.05%和2.45%（图1）。

图1

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图12018年良星77不同穗位粒位种子的活力指标比较

A：单株干重;B：活力指数。D2：花后2 d;D6：花后6 d;*表示差异达5%显著水平;**表示差异达1%显著水平。下同
Fig. 1Comparison of vigor index of seeds in different spike and grain positions of Liangxing 77 in 2018

A: Plant dry weight; B: Vigor index. D2: 2 days after anthesis; D6: 6 days after anthesis; * represents significant difference at the 5% level; ** represents significant differences at the 1% level. The same as below

2019年,对良星77不同穗位种子来说,花后6 d喷施DA-6可显著提高穗上部、穗中部种子的单株干重及活力指数,其中,对穗上部种子的单株干重及活力指数分别提高6.53%和4.40%,对穗中部种子的单株干重及活力指数分别提高6.18%和5.33%。花后2 d喷施DA-6使穗上部种子的单株干重和活力指数显著降低,与对照相比,分别显著降低4.90%和2.51%。对不同粒位的小麦种子来说,花后6 d喷施DA-6显著提高了下位粒、上位粒的单株干重及活力指数,其中,下位粒种子的单株干重及活力指数分别提高2.84%和2.38%,下位粒种子的单株干重及活力指数分别提高7.30%和4.06%;花后2 d喷施DA-6未见显著影响（图2）。

图2

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图22019年良星77不同穗位粒位种子的活力指标比较

A：单株干重;B：活力指数
Fig. 2Comparison of vigor index of seeds in different spike and grain positions of Liangxing 77 in 2019

A: Plant dry weight; B: Vigor index

同年,对山农23号来说,花后6 d喷施DA-6则使3个穗部位种子的单株干重与活力指数均显著提高。对穗上部种子的单株干重和活力指数分别显著提高8.44%和13.50%,对穗中部种子的单株干重和活力指数分别显著提高13.85%和11.55%,对穗下部种子的单株干重及活力指数则分别提高5.62%和5.24%。花后2 d喷施DA-6使穗上部、中部种子的活力显著降低,其中,穗上部种子的单株干重和活力指数分别极显著降低8.55%和10.52%,穗中部种子的单株干重和活力指数分别极显著降低5.78%和6.52%。对山农23号不同粒位的小麦种子来说,与良星77一样,花后6 d喷施DA-6显著提高了上位粒的单株干重与活力指数,分别提高5.75%和5.50%;花后2 d喷施DA-6未见显著影响（图3）。

图3

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图32019年山农23号不同穗位粒位种子的活力指标比较

A：单株干重;B：活力指数
Fig. 3Comparison of vigor index of seeds in different spike and grain positions of Shannong 23 in 2019

A: Plant dry weight; B: Vigor index

综上所述,花后6 d喷施DA-6可显著提高种子活力,并且种子活力的提高主要表现在穗上部、穗中部及上位粒的种子上,DA-6喷施有效降低了不同穗粒位种子之间的活力差异,使小麦不同穗位、粒位种子的活力整体均匀提高。

2.2 DA-6喷施对小麦籽粒发育进程千粒重和化学成分的影响

2018年良星77各处理组与对照组的千粒重及蛋白质含量均表现为随时间延长而上升（图4-A和图5）,结果表明,花后6 d喷施6 g·L^-1 DA-6的效果最好,与对照相比,千粒重、相对蛋白含量及绝对蛋白含量在花后40 d时分别提高了4.53%、4.79%和10.57%,同时在相对淀粉含量无显著变化的情况下绝对淀粉含量提高4.52%。而花后2 d喷施6 g·L^-1 DA-6则使千粒重、相对蛋白及绝对蛋白含量分别降低2.37%、6.15%和8.41%。

图4

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图4良星77、山农23号种子发育进程中千粒重变化

A：2018年良星77千粒重;B：2019年良星77千粒重;C：2019年山农23号千粒重。D2：花后2 d;D6：花后6 d;D25：花后25 d;D30：花后30 d;D35：花后35 d;D40：花后40 d。下同
Fig. 4Changes of 1000-grain weight during the seed development of Liangxing 77 and Shannong 23

A: 1000-grain weight of Liangxing 77 in 2018; B: 1000-grain weight of Liangxing 77 in 2019; C: 1000-grain weight of Shannong 23 in 2019. D2: 2 days after anthesis; D6: 6 days after anthesis; D25: 25 days after anthesis; D30: 30 days after anthesis; D35: 35 days after anthesis; D40: 40 days after anthesis. The same as below

图5

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图52018年良星77种子发育进程中化学成分变化

A：相对蛋白质含量;B：绝对蛋白质含量;C：相对淀粉含量;D：绝对淀粉含量
Fig. 5Chemical composition changes during seed development of Liangxing 77 in 2018

A: Relative content of protein, RCP; B: Absolute content of protein, ACP; C: Relative content of starch, RCS; D: Absolute content of starch, ACS

2019年良星77和山农23号的千粒重均表现为随着籽粒灌浆期的延长,千粒重先升高后降低;但相对蛋白及绝对蛋白含量则表现为随着时间的延长而渐升。同时,两品种均表现为花后6 d喷施6 g·L^-1的DA-6时千粒重及蛋白质含量最高。此处理下良星77花后40 d收获时,与对照相比,千粒重无显著差异,但相对蛋白质含量、绝对蛋白质含量分别显著提高2.42%和2.46%;淀粉含量略有下降但未达显著水平;而良星77花后2 d喷施DA-6的千粒重、相对蛋白、绝对蛋白含量、相对淀粉、绝对淀粉含量无显著差异（图4-A、图4-B、图5—图6）。山农23号花后6 d喷施6 g·L^-1 DA-6后千粒重、绝对蛋白含量均显著提高,提高幅度分别为2.28%和3.23%,淀粉含量无显著变化;花后2 d喷施DA-6对千粒重、相对蛋白质含量、绝对蛋白含量及淀粉含量无显著影响（图4-C、图7）。因此,两品种花后6 d喷施DA-6的小麦种子,最终收获时蛋白质含量显著高于对照。

图6

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图62019年良星77种子发育进程中化学成分变化

A：相对蛋白质含量;B：绝对蛋白质含量;C：相对淀粉含量;D：绝对淀粉含量
Fig. 6Chemical composition changes during seed development of Liangxing 77 in 2019

A: Relative content of protein, RCP; B: Absolute content of protein, ACP; C: Relative content of starch, RCS; D: Absolute content of starch, ACS

图7

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图72019年山农23号种子发育进程中化学成分变化

A：相对蛋白质含量;B：绝对蛋白质含量;C：相对淀粉含量;D：绝对淀粉含量
Fig. 7Chemical composition changes during seed development of Shannong 23 in 2019

A: Relative content of protein, RCP; B: Absolute content of protein, ACP; C: Relative content of starch, RCS; D: Absolute content of starch, ACS

2.3 DA-6喷施对种子发育过程中物质代谢酶基因表达的影响

为了验证DA-6喷施对种子发育过程中物质代谢酶基因表达的影响,选取前期转录组分析中喷施DA-6后显著上调的6个差异表达基因（茎秆、叶片和籽粒中各有2个）,这些基因分别与蛋白质、蔗糖代谢及抗逆性相关,对2019年花后6 d喷施6 g·L^-1 DA-6的处理组和对照组所取样品进行荧光定量PCR分析。

由图8可知,花后6 d喷施DA-6使得良星77和山农23号叶片中蛋白磷酸酶和蔗糖磷酸合成酶基因的表达量均极显著提高,其中良星77花后12 d所取叶片蛋白磷酸酶和蔗糖磷酸合成酶基因表达量提高幅度更大,而山农23号则表现为花后22 d所取叶片2个基因表达量大幅度提高。

图8

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图8DA-6喷施后小麦叶片蛋白磷酸酶和蔗糖磷酸合成酶基因表达

A：良星77叶片中蛋白磷酸酶基因;B：山农23号叶片中蛋白磷酸酶基因;C：良星77叶片中蔗糖磷酸合成酶基因;D：山农23号叶片中蔗糖磷酸合成酶基因
Fig. 8Gene expression of protein phosphatase and sucrose phosphate synthase in wheat leaves after DA-6 spraying

A: Protein phosphatase gene in Liangxing 77 leaves; B: Protein phosphatase gene in Shannong 23 leaves; C: Sucrose phosphate synthase gene in Liangxing 77 leaves; D: Sucrose phosphate synthase gene in Shannong 23 leaves

由图9可知,对茎秆中Ⅰ类hsp和hsp70表达量测定表明,在良星77中2个基因均表现为花后12 d表达量极显著提高,而花后22 d DA-6喷施处理与对照无显著差异;山农23号表现为花后22 d 2个基因表达量大幅度提高,而花后12 dⅠ类hsp表达量与对照相比也显著提高。

图9

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图9DA-6喷施对小麦茎秆Ⅰ类hsp和hsp70表达的影响

A：良星77茎秆中Ⅰ类hsp;B：山农23号茎秆中Ⅰ类hsp;C：良星77茎秆中hsp70;D：山农23号茎秆中hsp70
Fig. 9Effects of DA-6 spraying on gene expression of Class I hsp and hsp70 in wheat stems

A: Class I hsp gene in Liangxing 77 stems; B: Class I hsp gene in Shannong 23 stems; C: hsp70 gene in Liangxing 77 stems; D: hsp70 gene in Shannong 23 stems

籽粒中蛋白质二硫键异构酶和hsp82表达量测定表明,2个小麦品种在2个DA-6喷施后的取样期中均表现为显著提高,其中,以山农23号花后22 d蛋白质二硫键异构酶基因的提高幅度为最大;而对于hsp82,两品种均表现为花后22 d提高幅度最大（图10）。

图10

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图10DA-6喷施对小麦籽粒中蛋白质二硫键异构酶和hsp82表达的影响

A：良星77籽粒中蛋白质二硫键异构酶基因;B：山农23号籽粒中蛋白质二硫键异构酶基因;C：良星77籽粒中hsp82;D：山农23号籽粒中hsp82
Fig. 10Effects of DA-6 on the expression of protein disulfide isomerase and hsp82 in wheat grains

A: Protein disulfide isomerase gene in Liangxing 77 grains; B: Protein disulfide isomerase gene in Shannong 23 grains; C: hsp82 gene in Liangxing 77 grains; D: hsp82 gene in Shannong 23 grains

因此,良星77和山农23号DA-6喷施后叶片蛋白磷酸酶和蔗糖磷酸合成酶基因表达水平显著上调,茎秆中Ⅰ类hsp和hsp70表达量显著提高,籽粒中蛋白质二硫键异构酶和hsp82也显著上调,表明花后6 d喷施6 g·L^-1 DA-6可以显著促进蛋白、蔗糖代谢关键酶基因和热激蛋白基因的表达,有利于种子贮藏物质的合成,使其具有更强的适应性。

2.4 DA-6喷施对小麦种子萌发期代谢酶的影响

以良星77和山农23号2019年花后6 d喷施DA-6的样品（CK,6 g·L^-1）为材料,检测种子萌发期α-淀粉酶活性差异。

对山农23号萌发过程中8个时间点α-淀粉酶活性的检测表明,山农23号对照和处理组均表现为随着萌发时间的延长其α-淀粉酶活性会逐渐增强（图11-A）,大部分时间点的α-淀粉酶活性表现为喷施DA-6后的小麦种子酶活性高。由图可以看出,萌发72 h后,山农23号喷施DA-6的种子α-淀粉酶活性显著高于对照,与对照相比酶活性提高了15.67%。

图11

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图11DA-6喷施对良星77和山农23号α-淀粉酶活性的影响

A：山农23号;B：良星77;C：山农23号
Fig. 11Effect of DA-6 spraying on the α-amylase activity of Liangxing 77 and Shannong 23

A: Shannong 23; B: Liangxing 77; C: Shannong 23

2个品种种子萌发72 h后的α-淀粉酶活性比较结果表明,花后6 d喷施可显著提高α-淀粉酶活性,其中良星77比对照的α-淀粉酶活性提高了27.02%,山农23号只提高15.67%（图11-B、图11-C）。综上所述,对种子萌发期酶活性测定表明,喷施DA-6后的种子α-淀粉酶的活性显著提高,说明幼苗物质转化能力较强。

3 讨论

3.1 DA-6喷施对小麦不同穗粒位种子活力的影响

李孟良等^[34]、刘旭欢等^[35]研究表明小麦中穗下部种子活力高于穗上部的种子活力,且下位粒的活力要大于上位粒活力。本研究前期对小麦DA-6喷施的适宜时期和适宜浓度开展了连续2年的研究,表明DA-6适宜喷施浓度为6 g·L^-1,适宜喷施时期为花后6 d。本研究证明,花后适时喷施合适浓度的DA-6确实可通过提高单株干重来提高种子活力,即提高了幼苗的干物质积累。进一步通过不同穗位种子的发芽试验证明,DA-6对本来活力最高的穗下部种子其提高幅度不明显,但对活力相对较低的穗中部和穗上部种子提高幅度明显;而对不同粒位种子的发芽试验也说明,DA-6更能提高上位粒的种子活力。2年的试验结果略有差异,2018年度花后2 d喷施DA-6后使下位粒的种子活力降低,而2019年花后2 d喷施DA-6则未出现下位粒种子活力降低的现象;花后6 d喷施DA-6则均表现为提高上位粒种子活力,2年结果的差异究其原因可能为2年气候不同使结果出现波动,但对整体结果影响不大。另外2019年度加大喷施量,喷施更均匀,喷施效果更好。本研究表明,花后6 d喷施DA-6通过调节不同穗粒位种子活力,最终使收获的不同部位的种子间活力差异更小,使小麦不同穗位、粒位种子的活力整体均匀提高。

此外,有研究指出,萌发72 h的α-淀粉酶活性可以用来快速评价小麦种子活力的高低^[36]。本研究对2个小麦品种的种子萌发72 h后的α-淀粉酶活性比较表明,花后6 d DA-6喷施可显著提高种子萌发过程中的α-淀粉酶活性,表明DA-6促进了淀粉的分解利用和种胚萌动,为种子萌发提供物质与能量来源,从而促进种子的萌发,达到苗强苗壮的目的。

3.2 DA-6喷施对小麦生育过程中千粒重及化学成分的影响

任红等^[37]在玉米第8—10叶全展期2次喷施DA-6使玉米产量提高。杨学举^[38]研究发现对小麦增施氮肥可提高其蛋白质含量,降低直链淀粉含量,可改良小麦的淀粉特性。本研究中,花后6 d喷施DA-6使小麦从花后30 d开始千粒重及绝对蛋白含量、绝对淀粉含量比对照高,生长到花后35 d时提高幅度最大,而相对淀粉含量则成相反趋势,表明花后适宜时期喷施DA-6可提高千粒重,同时使蛋白质所占比例升高,使其营养价值提高,这与前人研究结果相符。

3.3 DA-6喷施提高小麦种子活力的机理

WEN等^[33]研究表明,山农23号在花后一周喷施4和6 g·L^-1 DA-6处理,籽粒千粒重和籽粒蛋白质含量均显著高于对照。花后24 d取样进行RNA-seq分析表明,6 g·L^-1 DA-6处理的样品中参与旗叶蔗糖合成、茎中内质网蛋白加工、种子中淀粉合成和内质网蛋白加工的差异表达基因均比对照有显著上调。本研究通过qRT-PCR分析,表明花后喷施DA-6调节了茎中内质网蛋白质合成、旗叶中蔗糖合成,籽粒中淀粉和蔗糖代谢,符合前人研究结果,DA-6使小麦在生育过程中籽粒的蛋白质含量升高,千粒重提高,最终使喷施DA-6后的小麦种子千粒重、蛋白质含量和种子活力提高。

4 结论

花后6 d喷施6 g·L^-1 DA-6可以促进蛋白、蔗糖代谢关键酶基因和热激蛋白基因的表达,有利于种子贮藏物质合成,使种子蛋白质含量升高,千粒重提高;显著提高收获种子穗上部、穗中部及上位粒的活力,降低了不同穗位粒位之间的种子活力差异,并使种子萌发过程中α-淀粉酶活性提高,物质转化能力提高,促进幼苗干物质积累,最终提高了种子活力。

参考文献原文顺序
文献年度倒序
文中引用次数倒序
被引期刊影响因子

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. Understanding the molecular pathways associated with seed vigor
Plant Physiology and Biochemistry, 2012,60:196-206.

DOI:10.1016/j.plaphy.2012.07.031 URL [本文引用: 1]

Farmers and growers are constantly looking for high quality seeds able to ensure uniform field establishment and increased production. Seed priming is used to induce pre-germinative metabolism and then enhance germination efficiency and crop yields. It has been hypothesized that priming treatments might also improve stress tolerance in germinating seeds, leaving a sort of 'stress memory'. However, the molecular bases of priming still need to be clarified and the identification of molecular indicators of seed vigor is nowadays a relevant goal for the basic and applied research in seed biology. It is generally acknowledged that enhanced seed vigor and successful priming depend on DNA repair mechanisms, activated during imbibition. The complexity of the networks of DNA damage control/repair functions has been only partially elucidated in plants and the specific literature that address seeds remains scanty. The DNA repair pathways hereby described (Nucleotide and Base Excision Repair, Non-Homologous End Joining, Homologous Recombination) play specific roles, all of them being critical to ensure genome stability. This review also focuses on some novel regulatory mechanisms of DNA repair (chromatin remodeling and small RNAs) while the possible use of telomere sequences as markers of aging in seed banks is discussed. The significant contribution provided by Electron Paramagnetic Resonance in elucidating the kinetics of seed aging, in terms of free radical profiles and membrane integrity is reported. (C) 2012 Elsevier Masson SAS.

[5]

ZHANG

, XIE

, XU

, JIANG

. Dissipation of the plant growth regulator hexanoic acid 2-(diethylamino) ethylester in pakchoi and soil
International Journal of Environmental and Analytical Chemistry, 2008,88(8):561-569.

DOI:10.1080/03067310802002490 URL [本文引用: 1]

[6]

JIANG

, JIANG

, HE

, ZHANG

, PAN

. Dissipation of diethyl aminoethyl hexanoate (DA-6) residues in pakchoi, cotton crops and soil
Bulletin of Environmental Contamination and Toxicology, 2012,88(4):533-537.

DOI:10.1007/s00128-012-0565-3 URL [本文引用: 1]

QuEChERS procedure and acetonitrile extraction, oscillation and ultrasonic procedure followed by GC-MS and LC-MS/MS (QqQ) detections were established for determination of diethyl aminoethyl hexanoate (DA-6) residues in pakchoi, cotton leaf, cotton seed and soil. At concentration levels of 0.005-1 mg kg(-1), recoveries were in the range of 80.5%-103.3%, with a RSD less than 14.2%. The LOQs of methods were 0.005, 0.003, 0.005 and 0.001 mg kg(-1) for the pakchoi, cotton leaf, soil and cotton seed samples, respectively. DA-6 was applied in supervised field trials at GAP conditions to pakchoi and cotton. It was found that the dissipation half-lives of DA-6 were 5.4-8.2 days and 1.1-2.2 days and 1.5-1.9 days in cotton crop, pakchoi and soil respectively. At harvest, no detectable residues (< LOD) were found in cotton samples. However, residues was detected in pakchoi (0.007-0.013 mg/kg) in Beijing and soil (0.008-0.014 mg/kg) in Changsha in 2008.

[7]

马庆华, 谭晓红, 梁丽松, 王贵禧. 生长调节剂DA-6和DCPTA处理对冬枣果实发育过程中活性氧及其相关生理指标的影响
食品科学, 2011,32(8):296-299.

URL [本文引用: 1]

The effects of the plant growth regulators DA-6 and DCPTA on the metabolism of reactive oxygen species in Ziziphus jujuba Mill. cv. Dongzao during growth were investigated. DA-6, DCPTA and the control (water) were sprayed on the tree crown 3 times at blossom, 20 days and 40 days after blossom, respectively. Fruit samples were collected regularly during different growing periods. In this study, the generation rate of superoxide anion free radicals, the contents of H2O2 and MDA and the activities of catalase (CAT), superoxide dismutase (SOD) and peroxidase (POD) were measured. The results showed that DA-6 and DCPTA treatments had no significant effect on changes in the generation rate of superoxide anion free radicals in ‘Dongzao’ jujube, but could accelerate its decline. DA-6 treatment increased H2O2 content significantly than DCPTA treatment. Both treatments dramatically increased the activities of SOD and POD in comparison with the control. DCPTA treatment increased the activity of CAT significantly at the end of the growth and inhibited the production of MDA, whereas DA-6 treatment had inhibitory effect on the activity of CAT. These finding can provide experimental data and theoretical supports for delaying the postharvest senescence of‘Dongzao’jujube.

Q H

, TAN

X H

, LIANG

L S

, WANG

G X

. Effects of plant growth regulators DA-6 and DCPTA on reactive oxygen species and related physiological indices of Ziziphus jujuba Mill. cv.‘Dongzao’during growth
Food Science, 2011,32(8):296-299. (in Chinese)

URL [本文引用: 1]

崔洪秋, 冯乃杰, 孙福东, 刘春娟, 何天明, 赵晶晶, 刘洋, 龚屾, 师臣, 郑殿峰. DTA-6和S₃₃₀₇对大豆存留荚和脱落荚生理调控的效应
中国农业科学, 2016,49(15):2921-2931.

DOI:10.3864/j.issn.0578-1752.2016.15.006 URL [本文引用: 1]

【Objective】The objective of this experiment is to study the effects of plant growth promoter and plant growth retardator on the difference of physiology in normal and abscission pods of soybean during the pods development in the semi-arid area, to discuss the physiological effects of plant growth regulator on reduction of the abscission of flowers and pods in soybean, and try to find the ways to increase soybean output. 【Method】 Diethyl aminoethyl hexanoate（DTA-6）, Uniconazole（S3307）and water were foliage sprayed at initial flowering (R1) stage of 3 varieties including Suinong 28 (SN28), Kenfeng 16 (KF16), and Hefeng 50 (HF50) in the Lindian county, Daqing city of Heilongjiang province in 2012 and 2013. The first time to take samples was 35 d (R5) after spraying, since then, the normal and abscission pods treated with CK and treatments were collected every seven days. The pod coats and seeds were separated, quick frozen for 30 min in the liquid nitrogen, and then stored in low temperature refrigerator(-40℃). Determination was carried out when all the samples were collected. The regulating differences of DTA-6 and S3307 between normal and abscission pods in oxygen free radical metabolism, related abscisic enzymes and soluble matter in soybean were determined and compared. 【Result】 The results demonstrated that during the pod development process, the malonaldehyde (MDA) content, superoxide dismutase (SOD) activity, peroxidase (POD) activity, soluble sugar and soluble protein of abscission pods were significantly higher than normal pods, the abscission cellulose (AC) activity, polygalacturonase (PG) activity in abscission pods were significantly lower than normal pods. DTA-6 and S3307 could regulate the physiology of normal and abscission pods in soybean. Although DTA-6 and S3307 showed different process in regulating effect, but both in the process had many similar results. The regulating effects of DTA-6 and S3307 showed as follows: DTA-6 and S3307 treatments decreased MDA content, improved SOD, POD activities in normal pods, and decreased AC and PG activities at pod-filling early stage in SN28 and KF16, but decreased at pod-filling late stage in HF50. The physiological difference of normal and abscission pods could change with the pods developmental process in soybean. The extents of increase and decrease were different during pods development process. The physiological difference was influenced by genetically controlled factors.【Conclusion】Compared with normal pods, the physiological indexes of abscission pods were improved, plasma membrane peroxidation was enhanced, balance of protective enzyme system was broken, soluble material were increased, related abscission enzymes activities reduced, and these were affected by environmental factors. The normal pods regulated by DTA-6 and S3307 showed a positive response to biological membrane damage, osmotic adjustment, protective enzyme system, related abscission enzymes activity reduced, especially in improving the physiological regulation and self-repairing ability.

CUI

H Q

, FENG

N J

, SUN

F D

, LIU

C J

, HE

T M

, ZHAO

J J

, LIU

, GONG

, SHI

, ZHENG

D F

. Effects of DTA-6 and S₃₃₀₇ on physiological regulation in normal and abscission pods of soybean
Scientia Agricultura Sinica, 2016,49(15):2921-2931. (in Chinese)

DOI:10.3864/j.issn.0578-1752.2016.15.006 URL [本文引用: 1]

刘春娟, 冯乃杰, 郑殿峰, 宫香伟, 孙福东, 石英, 崔洪秋, 张盼盼, 赵晶晶. 植物生长调节剂S₃₃₀₇和DTA-6 对大豆源库碳水化合物代谢及产量的影响
中国农业科学, 2016,49(4):657-666.

DOI:10.3864/j.issn.0578-1752.2016.04.005 URL [本文引用: 1]

【Objective】The aim of this paper was to investigate the effect of plant growth regulators on carbohydrate content, sucrose metabolizing enzyme activities and yield of soybean in the source-sink theory aiming to provide scientific basis for PGRs on agricultural production.【Method】Foliar spray field experiments were conducted successively in 2013 and 2014 in the main soybean producing zone, Heilongjiang Bayi Agricultural University. The experiment was in the early flowering by foliar spray one time. Taking Hefeng50 and Kefeng16 soybean varieties as the main research materials. Two chemical regulators, 2-N,N-diethylamino ethylcaproate (DTA-6, 60 mg·L-1) and uniconazole (S3307, 50 mg·L-1) were tested with water as the control. The first sampling was carried out after sprayed 30 d, and then leaf samples were collected once every five days. The contents of sucrose, starch or fructose in leaves and seeds, and invertase activity,SPS or SS activity in leaves were measured. The grain yield of soybean were investigated.【Result】At the early seed filling stage（30-35 days since spraying of DTA-6 and S3307）, the sucrose,fructose and starch contents of leaves showed the downtrends, and that of seeds showed increasing trends, which showed that more carbohydrates were applied to seeds growth and development. At the middle seed filling stage（35-45 days since spraying of DTA-6 and S3307）, the sucrose, fructose and starch contents of leaves always showed increasing trends by S3307 .The sucrose and fructose contents were all higher than the control, which was based to provide sufficient materials for the grain filling. At the late seed filling stage（after the 50th day since the spraying by DTA-6 and S3307）, the sucrose contents of leaves reached the maximum by DTA-6 and S3307, and reached significant levels to CK. The starch content of leaves sprayed with S3307 were higher than CK, and the fructose contents of leaves sprayed with DTA-6 were higher than CK. The sucrose contents of seeds were significantly increased by both S3307 and DTA-6 treatments. The fructose contents of two varieties seeds were increased by S3307, but that of Hefeng50 seed was decreased by DTA-6. The starch content of Hefeng50 seed but that were of Kefeng16 seed were increased by both S3307 and DTA-6 treatments, which showed that PGRs were different effect for soybean varieties. With sucrose contents were increased, SPS and SS activities were improved by S3307 and DTA-6. In most times, foliar spray of S3307 and DTA-6 could significantly decrease invertase activity of leaves to regulate carbohydrate metabolism in different varieties. Thereby, sucrose was neutral for carbohydrate metabolism. Balance of carbohydrate metabolism in source and sink by S3307 and DTA-6 to significantly increase yield. To controls（H-CK and K-CK）,the yield of PGRs (H-S, H-D and K-S, K-D) treatments were significantly increased by 20.07%, 14.57% and 10.54%, 10.95%. By the correlation analysis, The sucrose contents of the leaves were positively correlated with SPS, SS activities and starch contents of leaves and the sucrose, fructose and starch contents of seeds (0.893**, 0.888** and 0.981** or 0.918**, 0.832 and 0.810), or were negatively correlated with activities and invertase activities and fructose contents of leaves (-0.872 and -0.862).【Conclusion】The activity of SS and SPS which are of great importance in sucrose synthesis were increased and the activity of invertase was decreased by PGRs, the source-sink physiological metabolism of carbohydrates was regulated to increase yield by S3307 were higher than DTA-6.

LIU

C J

, FENG

N J

, ZHENG

D F

, GONG

X W

, SUN

F D

, SHI

, CUI

H Q

, ZHANG

P P

, ZHAO

J J

. Effects of plant growth regulators S₃₃₀₇ and DTA-6 on carbohydrate content and yield in soybean
Scientia Agricultura Sinica, 2016,49(4):657-666. (in Chinese)

DOI:10.3864/j.issn.0578-1752.2016.04.005 URL [本文引用: 1]

孙晓慧, 李成亮, 陈剑秋, 刘龙飞, 王秋双, 杨帆. 不同胺鲜酯 (DA-6) 浓度及施用方式对菠菜生长的影响
北方园艺, 2017(13):122-128.

[本文引用: 1]

SUN

X H

, LI

C L

, CHEN

J Q

, LIU

L F

, WANG

Q S

, YANG

. Effects of different DA-6 concentrations and application methods in growth of spinach
Northern Horticulture, 2017(13):122-128. (in Chinese)

[本文引用: 1]

[11]

周旭红, 梁华, 李纯佳, 蒋亚莲. 植物生长调节剂对香石竹生长发育的影响研究
中国农学通报, 2018,34(2):23-27.

[本文引用: 1]

ZHOU

X H

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C J

, JIANG

Y L

. Effect of compound sodium nitrophenolate and DA-6 on carnation growth
Chinese Agricultural Science Bulletin, 2018,34(2):23-27. (in Chinese)

[本文引用: 1]

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XIE

, GU

, MENG

, LI

, WANG

, QU

, WEI

. Exogenous DCPTA ameliorates simulated drought conditions by improving the growth and photosynthetic capacity of maize seedlings
Scientific Reports, 2017,7(1):1-13.

DOI:10.1038/s41598-016-0028-x URL [本文引用: 1]

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张志芳, 贾海丽, 张小会. DA-6对番茄生长的影响
现代农业科技, 2012,6:193-194+197.

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ZHANG

Z F

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H L

, ZHANG

X H

. Influence of DA-6 for tomato growth
Modern Agricultural Science and Technology, 2012,6:193-194+197. (in Chinese)

[本文引用: 1]

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陈艳丽, 范飞, 王旭, 李绍鹏, 曹振木. DA-6对高温胁迫下黄灯笼辣椒幼苗的影响
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CHEN

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Z M

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[本文引用: 1]

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王丹, 马青松, 范佐旺, 李小婷, 宛骏, 袁蕾, 陈晓鹭, 邹纯礼, 庞玉新. DA-6 对冬季迟缓期的艾纳香生长和有效成分含量的影响
中国现代中药, 2015,17(11):1184-1187+1192.

[本文引用: 1]

WANG

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Modern Chinese Medicine, 2015,17(11):1184-1187+1192. (in Chinese)

[本文引用: 1]

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杨修一, 李圣会, 梅宇超, 杨广, 孙晓慧, 赵晨浩, 张民, 李成亮. DA-6对水培生菜生长及生理特性的影响
农业环境科学学报, 2017,36(1):32-38.

[本文引用: 1]

YANG

X Y

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S H

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. Effects of DA-6 on growth and physiological characteristics in hydroponic lettuce
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[本文引用: 1]

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聂乐兴, 姜兴印, 吴淑华, 王燕, 李俊虎, 戈大庆, 张吉旺, 刘鹏. 胺鲜酯对高产夏玉米产量及叶片光合羧化酶和保护酶活性的影响
应用生态学报, 2010,21(10):2558-2564.

URL [本文引用: 1]

A field plot experiment was conducted to study the effects of foliar spraying DA-6 at the rates of 10, 20, and 40 mg·L-1at jointing stage on the leaf photosynthetic carboxylase and protective enzyme activities and grain yield of high-yielding summer maize cultivar Denghai 661. Comparing with the control (foliar spraying surfactant and water), spraying 10, 20, and 40 mg·L-1 of DA-6 increased the grain yield of Denghai 661 significantly, with the increment being 10.0%, 8.9%, and 9.4%, respectively, but no significant differences were observed among the DA-6 treatments. Different concentration DA-6 increased the leaf area index, net photosynthetic rate, and RuBPCase and PEPCase activities significantly, and the promotion effects on the net photosynthetic rate and RuBPCase and PEPCase activities increased with increasing concentration of DA-6. After treated with DA-6, the leaf superoxide dismutase, catalase, peroxidase, and gultathione S-transferase activities and the leaf soluble protein content at the stages of silking, grain-filling, milky, and wax maturity all increased significantly, and the leaf malondialdehyde decreased significantly, compared with the control. The catalase activity increased with increasing DA-6 concentration, but the other indices had no significant differences among the DA-6 treatments.

NIE

L X

, JIANG

X Y

, WU

S H

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, LI

J H

, GE

D Q

, ZHANG

J W

, LIU

. Effects of DA-6 on leaf photosynthetic carboxylase and protective enzyme activities and grain yield of high-yielding summer maize
Chinese Journal of Applied Ecology, 2010,21(10):2558-2564. (in Chinese)

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聂乐兴, 姜兴印, 吴淑华, 张吉旺, 刘鹏. 胺鲜酯对高产玉米的调控作用研究
玉米科学, 2010,18(6):33-37.

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Journal of Maize Sciences, 2010,18(6):33-37. (in Chinese)

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ZHOU

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. DA-6 promotes germination and seedling establishment from aged soybean seeds by mediating fatty acid metabolism and glycometabolism
Journal of Experimental Botany, 2019,70(1):101-114.

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