化学调控和氮肥对高密度下春玉米光热水利用效率和产量的影响

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刘笑鸣^,¹, 顾万荣^,¹, 李从锋², 张立国³, 王明泉³, 龚士琛³, 陈喜昌³, 李彩凤¹, 魏湜¹, 李文华³

¹东北农业大学农学院,哈尔滨 150030

²中国农业科学院作物科学研究所,北京 100081

³黑龙江省农业科学院玉米研究所,哈尔滨 150086

Effects of Chemical Regulation and Nitrogen Fertilizer on Radiation, Heat and Water Utilization Efficiency and Yield of Spring Maize Under Dense Planting Condition

LIU XiaoMing^,¹, GU WanRong^,¹, LI CongFeng², ZHANG LiGuo³, WANG MingQuan³, GONG ShiChen³, CHEN XiChang³, LI CaiFeng¹, WEI Shi¹, LI WenHua³

¹College of Agronomy, Northeast Agricultural University, Harbin 150030

²Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, Beijing 100081

³Maize Research Institute, Heilongjiang Academy of Agricultural Sciences, Harbin 150086

通讯作者: 顾万荣,E-mail： wanronggu@163.com

责任编辑: 杨鑫浩
收稿日期:2020-04-8接受日期:2020-07-6网络出版日期:2020-08-01

基金资助:

国家重点研发计划课题.2016YFD0300103
国家重点研发计划课题.2017YFD0300506
国家重点研发计划项目黑龙江省配套资金项目.GX18B029
东北农业大学“学术骨干”基金.17XG23

Received:2020-04-8Accepted:2020-07-6Online:2020-08-01
作者简介 About authors
刘笑鸣,E-mail： liuxming1995@163.com。

摘要
【目的】探讨高密度下化学调控和氮肥对玉米光合特性、籽粒灌浆及光热水利用效率的影响,为玉米密植抗逆高产高效栽培提供依据。【方法】于2017—2018年哈尔滨区域玉米生长季,在大田高密度种植下（90 000株/hm²）,设置3个不同氮肥水平N100（100 kg·hm^-2）、N200（200 kg·hm^-2）和N300（300 kg·hm^-2）,7叶期喷施化控剂（玉黄金,30%胺鲜酯·乙烯利水剂）,研究化控和氮肥对高密度种植下玉米生长发育和光热水利用效率的影响。【结果】随氮肥施用量的增加,玉米叶片净光合速率（P_n）、最大光化学效率（F_v/F_m）、籽粒内源激素含量、灌浆速率、光热水利用效率（RUE、HUE和WUE）及产量均呈先升高后降低趋势,在施氮量200 kg·hm^-2下达到最大。与单施氮肥处理相比,化控和氮肥共同作用显著提高了叶片P_n和F_v/F_m,提高了籽粒内源激素含量和灌浆速率,RUE、HUE和WUE显著提高,使产量得到进一步增加。相关分析表明,灌浆速率与籽粒内源激素生长素（IAA）、细胞分裂素（CTK）、赤霉素（GA）和脱落酸（ABA）含量极显著正相关,产量与RUE、HUE和WUE显著正相关。【结论】高种植密度下,200 kg·hm^-2施氮量和化控显著改善玉米的光合特性,促进了籽粒灌浆进程,提高了光热水利用效率,显著提高了产量。
关键词： 玉米;光合特性;化学调控;氮肥;产量;资源利用效率

Abstract

【Objective】 This experiment was conducted to study the effects of chemical regulation and nitrogen fertilizer on photosynthesis, grain filling characteristics and radiation, heat and water utilization efficiency of maize under high planting density, so as to provide a theoretical basis for stress resistance, high yield and high efficiency of maize under high planting density. 【Method】 The experiment was performed with three nitrogen rates, N100 (100 kg·hm^-2), N200 (200 kg·hm^-2) and N300 (300 kg·hm^-2) under high planting density (90 000 plants/hm²) during the maize growth season from 2017 to 2018. Chemical control (Yuhuangjin, 30% amine fresh ester·ethyl hydroxide) was sprayed at the seventh leaf stage. The effects of chemical control and nitrogen application on maize growth and development and radiation (RUE), heat utilization efficiency (HUE) and water utilization efficiency (WUE) under high density were studied. 【Result】 With the nitrogen application increasing, the 8 indexes, including net photosynthetic rate (P_n), max photochemical efficiency (F_v/F_m) of maize leaves, the content of grain endogenous hormones, grain filling rate, radiation utilization efficiency, heat utilization efficiency, water utilization efficiency and yield, first increased and then decreased, and all reached the maximum at 200 kg·hm^-2 nitrogen rate. Compared with the single application of nitrogen fertilizer, the combination of chemical control and nitrogen fertilizer significantly increased P_n and F_v/F_m in leaves, endogenous hormone content in grains, grain filling rate, RUE, HUE, and WUE, which further increased grain yield. The correlation analysis showed that the grain filling rate was positively correlated with indole-3-acetic acid (IAA), cytokinin (CTK), gibberellin (GA) and abscisic acid (ABA), while the yield was positively correlated with RUE, HUE and WUE.【Conclusion】 Under high planting density, 200 kg·hm^-2 nitrogen application and chemical control significantly improved the photosynthetic characteristics, promoted the grain filling process, improved radiation, heat and water utilization efficiency, and significantly increased the yield.

Keywords：maize;photosynthetic characteristics;chemical regulation;nitrogen fertilizer;yield;resource utilization efficiency

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本文引用格式
刘笑鸣, 顾万荣, 李从锋, 张立国, 王明泉, 龚士琛, 陈喜昌, 李彩凤, 魏湜, 李文华. 化学调控和氮肥对高密度下春玉米光热水利用效率和产量的影响[J]. 中国农业科学, 2020, 53(15): 3083-3094 doi:10.3864/j.issn.0578-1752.2020.15.009
LIU XiaoMing, GU WanRong, LI CongFeng, ZHANG LiGuo, WANG MingQuan, GONG ShiChen, CHEN XiChang, LI CaiFeng, WEI Shi, LI WenHua. Effects of Chemical Regulation and Nitrogen Fertilizer on Radiation, Heat and Water Utilization Efficiency and Yield of Spring Maize Under Dense Planting Condition[J]. Scientia Acricultura Sinica, 2020, 53(15): 3083-3094 doi:10.3864/j.issn.0578-1752.2020.15.009

0 引言

【研究意义】黑龙江省是我国主要的玉米生产基地^[1],2018年玉米种植面积达到631.78×10⁴ hm²,总产达3 982.2×10⁴t。该地区土壤肥沃,光水资源丰富,有利于玉米的生长;但随着气候的波动,增加了极端气候事件的发生强度和频率,玉米生育前期温度较低且易发生春旱,生殖生长期易遭遇干旱或阴雨寡照,不利于玉米的生长发育和籽粒的形成,对产量造成巨大损失^[2,3,4,5]。增加种植密度是获得玉米高产的重要措施,有利于提高光、热和养分等资源利用率,依靠群体发挥增产潜力^[6,7]。但密度过大易导致玉米群体密闭,叶片互相遮阴,从而影响光合作用,并且增加了倒伏的风险,导致玉米产量降低^[8,9]。前人研究表明,探索不同生态区玉米产量潜力及突破技术途径,提高单产和资源利用效率是玉米栽培的重要研究方向^[10,11,12]。因此,通过调整栽培措施改善高密度下玉米生长发育和光热水利用效率,对提高玉米群体生产力和实现玉米可持续生产有重要意义。【前人研究进展】氮肥的合理运筹是当前农业生产中作物管理的重点。作为玉米需求量最大的营养元素,适量施氮有利于调控作物生长发育,改善光合性能,实现优质高产^[13]。氮素是叶绿素的重要组成元素,其含量显著影响叶片的叶绿素含量和光合生理特性^[14]。在籽粒灌浆期,低氮条件下玉米穗位叶的光合速率、气孔导度、表观量子产量等均低于高氮处理^[15]。陈晓璐等^[16]研究表明,合理增施氮肥可增强光系统性能,提高净光合速率,延缓灌浆期叶绿素含量降低和叶片衰老,为玉米高产提供基础。研究表明,群体产量的提高与资源的高效利用密不可分,密植条件下资源利用率的提高有利于充分发挥玉米的产量潜力^[17]。MOMEN等^[18]通过对不同氮肥水平下玉米产量及资源利用效率进行研究,发现不同氮肥用量对光能利用效率（RUE）、水分利用效率（WUE）和产量均有影响,其中,玉米产量主要由RUE决定。AGGARWAL等^[19]研究表明,随着施氮量的增加,作物生长日积累量和WUE显著增加,有利于产量的提高。而田间实际生产中,常因氮肥施用不合理导致群体光合性能降低^[20,21],养分利用效率和产量下降^[22]。近年来,作物化学调控技术已成为我国粮食高产高效栽培的重要组成部分。前人研究表明,喷施化控剂可提高玉米叶源的活性,保持较高的叶绿素含量和最大光化学效率,延长叶片光合功能持续期,有利于物质的积累和产量的提高^[23,24]。籽粒灌浆是营养物质不断向籽粒输送并积累的过程,是决定籽粒产量的重要阶段^[25,26]。化控处理可改变籽粒内源激素含量,调控籽粒发育和籽粒同化物代谢,显著提高了灌浆速率和灌浆持续时间,最终提高了粒重和产量^[27,28]。合理的化控技术是提高资源利用效率、实现作物高产的有效途径^[29]。化学调控提高了作物WUE和RUE,提高了光合作用和生物量,有利于产量的提高^[30,31]。【本研究切入点】施氮对玉米生长发育及化控剂对玉米调控作用的研究已有报道,但化学调控和氮肥共同作用对玉米叶片光合、籽粒激素、灌浆特性及光热水利用效率影响的研究较少。【拟解决的关键问题】本研究通过设置不同氮肥水平,并结合喷施化控剂处理,比较产量、叶片光合指标、籽粒内源激素含量、籽粒灌浆特性、资源利用效率等变化,研究高密度下化控与氮肥对春玉米产量形成及光热水利用效率的调控机制,旨在为黑龙江省高密度下玉米高产高效栽培提供理论依据。

1 材料与方法

1.1 试验设计

试验于2017年和2018年在哈尔滨市东北农业大学向阳试验基地（126°54′56.37″E,45°46′01.27″N）进行。土壤为黑钙土,土壤有机质25.25 g·kg^-1、全氮1.7 g·kg^-1、碱解氮118.21 mg·kg^-1、速效磷65.34 mg·kg^-1、速效钾179.35 mg·kg^-1,pH 6.85。玉米生长周期内气象数据由哈尔滨市农业科学院提供（表1）。

Table 1
表1
表12017年和2018年玉米生长季降雨量和平均温度
Table 1Daily precipitation and mean temperature during the maize growing season in 2017 and 2018

月份 Month	平均温度 Average temperature (℃)		降雨量 Precipitation (mm)		日照时数 Sunshine hours (h)		平均风速 Average wind speed (m·s^-1)
月份 Month	2017	2018	2017	2018	2017	2018	2017	2018
5月 May	16.7	16.6	51.8	12.2	261	257	4.68	3.14
6月 June	19.9	21.2	92.2	186.4	218	202	2.82	3.22
7月 July	24.7	24.9	50.2	172.1	252	159	2.62	2.95
8月 August	22.1	21.4	215.2	121.9	168	184	3.05	2.33
9月 September	15.0	15.3	37.0	76.7	200	178	2.81	2.56

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选用玉米品种“龙育365”为试验材料,2年均于4月30日播种,种植密度为90 000株/hm²,每穴播3粒,于3叶期间苗,9月25日收获测产。供试化控剂为玉黄金（30%胺鲜酯·乙烯利水剂,福建浩伦生物工程技术有限公司）,在玉米7展叶期,于上午7：00用喷雾器叶面均匀喷施玉黄金（Y）（每公顷375 mL,兑水450 kg）,对照喷施清水（CK）;设置3个氮肥施用量水平N100（100 kg·hm^-2）、N200（200 kg·hm^-2）和N300（300 kg·hm^-2）,种肥和玉米封垄前追肥以1:1比例施入尿素。磷钾肥分别施用100 kg·hm^-2磷酸二氢铵和100 kg·hm^-2硫酸钾,作为种肥一次施入。本试验采用裂区设计,主区设置化控剂玉黄金处理和清水处理,副区设置3个氮肥水平,共6个处理,3次重复,每小区8行,行长8 m,行距0.65 m,小区面积41.6 m²。其他农艺措施参照当地高产栽培措施。

1.2 测定项目及方法

1.2.1 光合指标每处理选取3株长势一致的代表性植株,于拔节期（7月5日）取玉米倒3叶,抽雄期（7月25日）、灌浆初期（8月5日）、乳熟期（8月25日）和完熟期（9月25日）取穗位叶,用 Li-6400便携式光合仪（Li-COR,美国）测定净光合速率（P_n）;用 FMS2 脉冲调制式荧光仪（Hansatech,英国）测定叶绿素荧光参数,叶片暗适应30 min后,测得最小荧光（F_o）和最大荧光（F_m）,PSⅡ最大光化学效率F_v/F_m = (F_m-F_o)/F_m。

1.2.2 籽粒激素及灌浆特性在玉米吐丝期,选取长势一致的植株挂牌标记。于开花后10、15、20、25、30 d每处理取3个果穗,剥取穗中部籽粒,液氮冷冻 30 min,在-40℃冰箱中保存。采用酶联免疫吸附法（ELISA）测定生长素（IAA）、赤霉素（GA）、细胞分裂素（CTK）和脱落酸（ABA）的含量。于开花后 10、15、20、25、30、40、45、50 d 取样,105℃杀青30 min,80℃烘干,测籽粒干重。参照张莉等^[32]方法用Logistic 方程对籽粒灌浆过程进行拟合并计算灌浆速率。

1.2.3 光热水利用效率光能利用效率（RUE）（%）= W×H/∑Q×100%,H为每克干物质燃烧时释放出的热量,玉米干重热值为1.807×10⁴ J·g^-1,W是测定期间干物质增加量（g）,∑Q是同期总光照辐射量（MJ·m^-2·d^-1）;热量利用效率（HUE）（kg·hm^-2·℃^-1·d^-1）= Y/∑t, Y为籽粒产量（kg·hm^-2）,∑t为生育期间≥10℃的有效积温（℃·d）;水分利用效率（WUE）（kg·hm^-2·mm^-1）= Y/ET, Y为籽粒产量（kg·hm^-2）,ET为生育期内总耗水量（mm）。

1.2.4 产量完熟期在每小区中间行选取13 m²（4行×5 m行长）的所有玉米进行收获,自然风干后于室内考种并计产（按14%标准含水量折算产量）。

1.3 数据分析

采用Microsoft 2007整理数据和作图,通过 Curve Expert 1.3 软件进行灌浆动态拟合,使用SPSS 18.0分析数据。

2 结果

2.1 高密度下化学调控和氮肥处理对叶片净光合速率（P_n）及最大光化学效率（F_v/F_m）的影响

由图1可知,叶片P_n和F_v/F_m在2年中随玉米的生长发育呈先升高后降低的趋势,均在灌浆初期达到最大值。随着施氮量的增加,P_n和F_v/F_m先升高后降低,表现为N200>N300>N100,其中P_n在N200下分别比N100和N300高21.9%—22.4%和7.9%—9.4%,F_v/F_m在N200下分别比N100和N300高17.2%—22.8%和4.4%—5.5%。化控处理显著提高了各氮肥水平下叶片P_n和F_v/F_m,在2017—2018年分别提高9.4%—14.5%和5.1%—9.3%。在化控和氮肥共同作用下,N200+Y处理下的P_n和F_v/F_m最高。

图1

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图1高密度下化学调控对叶片净光合速率（P_n）及最大光化学效率（F_v/F_m）的影响

N100+CK、N200+CK和N300+CK分别表示施氮量100、200和300 kg·hm^-2下喷施清水,N100+Y、N200+Y和N300+Y分别表示施氮量100、200和300 kg·hm^-2下喷施化控。下同
Fig. 1Effects of chemical regulation on net photosynthetic rate (P_n) and maximal photochemistry efficiency (F_v/F_m) in leaves under high density

N100+CK indicates 100 kg N·hm^-2+water treatment; N200+CK indicates 200 kg N·hm^-2+water treatment; N300+CK indicates 300 kg N·hm^-2+water treatment; N100+Y indicates 100 kg N·hm^-2+Yuhuangjin treatment; N200+Y indicates 200 kg N·hm^-2+Yuhuangjin treatment; N300+Y indicates 300 kg N·hm^-2+ Yuhuangjin treatment. The same as below

2.2 高密度下化学调控和氮肥处理对花后玉米籽粒激素含量的影响

由图2可以看出,籽粒生长素（IAA）和细胞分裂素（CTK）含量在灌浆过程中呈先升高后降低的趋势,均于吐丝后25 d达到最大值。随着施氮量的增加,IAA和CTK含量先升高后降低,化控处理显著提高了各氮肥水平下的籽粒IAA和CTK含量,在化控和氮肥共同作用下,N200+Y 处理的籽粒IAA和CTK含量最高。籽粒赤霉素（GA）含量在灌浆过程中呈降低趋势,随着施氮量的增加,GA含量先升高后降低,化控处理显著提高了各氮肥水平下籽粒GA含量,在化控和氮肥共同作用下,N200+Y处理籽粒GA含量最高。籽粒脱落酸（ABA）含量在灌浆过程中先升高后降低,在吐丝后25 d达到最大值。不同氮肥处理下,N200的ABA含量最高,化控处理显著提高了各氮肥水平下籽粒ABA含量,在化控和氮肥共同作用下,N200+Y处理的籽粒ABA含量最高。

图2

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图2高密度下化学调控对花后玉米籽粒激素含量的影响

Fig. 2Effects of chemical regulation on hormones content in maize grain after anthesis under high density

2.3 高密度下化学调控和氮肥处理对籽粒灌浆参数的影响

以开花后天数为自变量,开花后每隔 5 d 测得的百粒重为因变量,用Logistics方程模拟籽粒灌浆过程。由表2可以看出,随着施氮量的增加,最大灌浆速率和平均灌浆速率先升高后降低,在N200达到最大,达到最大灌浆速率时的天数和灌浆活跃期略有增加。施用化控剂减少了各氮肥水平下籽粒达到最大灌浆速率时的天数（T_max）,更快地达到最大灌浆速率,并提高了最大灌浆速率（V_max）和平均灌浆速率（V_m）,而对灌浆活跃期（P）影响不显著。在化控和氮肥共同作用下,N200+Y处理的籽粒灌浆速率和灌浆活跃期最大且达到最大灌浆速率时的天数最短。

Table 2
表2
表2高密度下化学调控对籽粒灌浆参数的影响
Table 2Effects of chemical regulation on the characteristic parameters of grain filling under high density

年份 Year	处理 Treatment	方程参数 Parameter of equation			籽粒灌浆参数 Parameter of grain filling
年份 Year	处理 Treatment	A	B	C	T_max (d)	V_max (g·100-kernel^-1·d^-1)	V_m (g·100-kernel^-1·d^-1)	P (d)
2017	N100+CK	32.30	47.86	0.1408	27.47	1.1370	0.5374	42.61
	N200+CK	33.84	44.48	0.1412	26.88	1.1946	0.5695	42.49
	N300+CK	31.12	53.52	0.1469	27.09	1.1429	0.5331	40.84
	N100+Y	33.09	41.39	0.1395	26.69	1.1540	0.5549	43.01
	N200+Y	36.39	40.18	0.1382	26.72	1.2573	0.6068	43.42
	N300+Y	32.89	40.84	0.1393	26.63	1.1454	0.5517	43.07
2018	N100+CK	30.61	60.83	0.1427	28.79	1.0920	0.5019	42.05
	N200+CK	34.16	53.91	0.1369	29.13	1.1691	0.5449	43.83
	N300+CK	34.50	51.45	0.1335	29.52	1.1514	0.5396	44.94
	N100+Y	32.75	54.24	0.1405	28.42	1.1503	0.5358	42.70
	N200+Y	37.00	42.72	0.1335	28.12	1.2349	0.5916	44.94
	N300+Y	34.64	46.71	0.1367	28.12	1.1838	0.5611	43.89

A：终极生长量;B：初值参数;C：生长速率参数;T_max：达到最大灌浆速率的天数;V_max：最大灌浆速率;V_m：平均灌浆速率;P：灌浆活跃期
A: The final grain weight; B: Initial parameter; C: Growth rate parameter; T_max: The time reaching the maximum grain-filling rate; V_max: Maximum grain-filling rate; V_m: Mean grain-filling rate; P: Active grain-filling period

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2.4 籽粒灌浆速率与激素含量的相关性分析

利用Logistic方程模拟玉米籽粒灌浆过程,结合表2各籽粒灌浆参数,计算出花后不同阶段玉米籽粒在不同处理下的灌浆速率,与同一时期籽粒内源激素含量进行相关分析（表3）。在2017年,玉米籽粒灌浆速率在花后10—20 d与IAA、CTK和ABA均呈显著正相关;2018年,籽粒灌浆速率在花后10—30 d与IAA含量呈显著正相关,在花后10—25 d与CTK含量呈显著正相关,10—20 d与GA含量呈显著正相关,在花后10、15和25 d与ABA含量呈显著正相关。表明在灌浆前期和中期,其灌浆速率与IAA、CTK、GA和ABA密切相关,IAA、CTK、GA和ABA的含量越高,其灌浆速率就越快。

Table 3
表3
表3籽粒灌浆速率与激素含量的相关性分析
Table 3Correlation coefficients of grain filling rate with grain hormones content

年份 Year	项目 Item	IAA	CTK	GA	ABA
2017	花后10d灌浆速率 Grain filling rate on 10 d after anthesis	0.95**	0.90*	0.79	0.88*
	花后15d灌浆速率 Grain filling rate on 15 d after anthesis	0.97**	0.92**	0.79	0.92**
	花后20d灌浆速率 Grain filling rate on 20 d after anthesis	0.98**	0.89*	0.78	0.91*
	花后25d灌浆速率 Grain filling rate on 25 d after anthesis	0.74	0.80	0.54	0.65
	花后30d灌浆速率 Grain filling rate on 30 d after anthesis	0.54	0.46	0.30	0.46
2018	花后10d灌浆速率 Grain filling rate on 10 d after anthesis	0.83*	0.89*	0.95**	0.95**
	花后15d灌浆速率 Grain filling rate on 15 d after anthesis	0.90*	0.97**	0.93**	0.91*
	花后20d灌浆速率 Grain filling rate on 20 d after anthesis	0.92**	0.90*	0.91*	0.68
	花后25d灌浆速率 Grain filling rate on 25 d after anthesis	0.88*	0.93**	0.70	0.89*
	花后30d灌浆速率 Grain filling rate on 30 d after anthesis	0.81*	0.41	0.41	0.28

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2.5 高密度下化学调控和氮肥处理对玉米光热水利用效率的影响

由表4可知,随施氮量的增加,2017年玉米光能利用效率（RUE）逐渐增加,2018年RUE呈先增加后降低趋势。2017—2018年热量利用效率（HUE）和水分利用效率（WUE）均随施氮量增加而先增加后降低,N200最大。化控显著增加了2017—2018年各氮肥水平下玉米的RUE、HUE和WUE,分别提高了2.2%—7.0%、8.7—10.1%和8.7%—10.0%。在化控和氮肥共同作用下,N200+Y处理的玉米RUE、HUE和WUE最大,光热水利用效率最高。

Table 4
表4
表4高密度下化学调控对玉米光热水利用效率的影响
Table 4Effect of chemical regulation on radiation, heat and water utilization efficiency under high density

处理 Treatment	光能利用效率 RUE (%)		热量利用效率HUE (kg·hm^-2·℃^-1·d^-1)		水分利用效率 WUE (kg·hm^-2·mm^-1)
处理 Treatment	2017	2018	2017	2018	2017	2018
N100+CK	2.42bc	2.43d	3.48b	2.95c	20.55c	19.14c
N200+CK	2.48b	2.57c	3.73ab	3.34ab	22.14abc	21.07bc
N300+CK	2.68a	2.47d	3.61b	3.05bc	21.44bc	19.33c
N100+Y	2.40c	2.62bc	3.69b	3.20bc	22.56abc	19.90bc
N200+Y	2.66a	2.70a	3.99a	3.65a	24.30a	24.03a
N300+Y	2.69a	2.67ab	3.71ab	3.34ab	23.22ab	22.03ab

同一列中不同小写字母表示差异显著（P<0.05）。下同
Different lowercase letters in the same column indicate significant difference (P<0.05). The same as below

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2.6 高密度下化学调控和氮肥处理对产量及其构成因素的影响

由表5可知,2017年和2018年不同处理下玉米产量基本一致。随着施氮量的增加,产量先升高后降低,N200达到最大,化控显著提高了各施氮量下的产量。从产量构成因素来看,化控和氮肥对穗数影响不大,而对穗粒数和千粒重有显著影响,穗粒数和千粒重均随施氮量的增加而先升高后降低,化控显著增加了各施氮量下的穗粒数和千粒重。所有处理中,N200+Y处理穗粒数和千粒重最大、产量最高,2017年和2018年产量分别达到12 646 kg·hm^-2和11 704 kg·hm^-2。

Table 5
表5
表5高密度下化控和氮肥对玉米产量及其构成因素的影响
Table 5Effect of chemical control and nitrogen fertilizers on maize yield and its components under high density

处理 Treatment	穗数 Spike (ear/hm²)		穗粒数 Kernel per ear		千粒重 1000-grain weight (g)		产量 Yield (kg·hm^-2)
处理 Treatment	2017	2018	2017	2018	2017	2018	2017	2018
N100+CK	81078a	80325a	541c	531c	332b	294c	10511c	9840bc
N200+CK	81654a	80793a	568b	550bc	327b	298bc	11548b	10430b
N300+CK	81782a	78685b	560b	533c	316c	298bc	11053bc	9204c
N100+Y	81657a	81052a	571b	556abc	340ab	306bc	11427b	9990bc
N200+Y	81683a	81184a	591a	581a	351a	327a	12646a	11704a
N300+Y	82150a	81167a	570b	566ab	339ab	314ab	11921b	10732ab

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2.7 产量与光合及效率的协调性比较

探讨2年中高密度下化学调控和氮肥处理对玉米叶片光合作用和群体光热水利用效率的调控效应,相关分析结果表明（图3）,除2018年F_v/F_m和2017年RUE与产量相关性不显著外,2年中P_n、F_v/F_m、RUE、HUE和WUE均与产量呈显著或极显著正相关关系。说明高密度下通过提高叶片光合及荧光特性,提高群体光、热、水利用效率,可实现产量的提高。

图3

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图3产量与群体光热水利用效率的相关性分析

Fig. 3Correlation analysis of yield with radiation, heat and water utilization efficiency of population

3 讨论

3.1 高密度下化学调控和氮肥处理对玉米光合作用的影响

光合作用是作物生长发育和产量形成的生理基础,光合能力的提高对增加籽粒产量有重要作用^[33]。净光合速率和最大光化学效率是重要的光合及荧光参数,反映了叶片的光合效率和光合作用强度。魏廷邦等^[34]研究认为,增施氮肥可增强玉米密植条件下的光合作用,提高叶片净光合速率。化控处理可以提高叶片叶绿素含量和净光合速率,使叶片维持较高的光合能力^[35]。前人研究表明,化控可以提高玉米净光合速率和抗氧化酶活性,延缓叶片衰老,并使光合生理功能得到改善^[36,37]。本研究表明,高密度下各氮肥处理中,随着施氮量的提高,叶片P_n和F_v/F_m先升高后降低,施氮量200 kg·hm^-2下显著提高了叶片光合能力。与单施氮肥处理相比,在氮肥和化学调控共同作用下,叶片P_n和F_v/F_m显著提高,其中,200 kg·hm^-2施氮量下施用化控时光合及荧光参数达到最大。说明化学调控能够在氮肥促进叶片光合生理特性的基础上,进一步增强叶片对光能的利用,提高了光合能力。相关分析表明,产量与P_n和F_v/F_m显著正相关,说明在施氮量200 kg·hm^-2下施用化控剂通过增强叶片P_n和F_v/F_m,有利于最终玉米产量的提高。

3.2 高密度下化学调控和氮肥处理对籽粒灌浆及其激素调控的影响

籽粒灌浆是玉米产量形成过程中重要的生育阶段,灌浆时间和灌浆速率决定了玉米粒重和产量的形成^[26]。申丽霞等^[38]研究表明,在高密度栽培条件下氮肥可有效调控籽粒灌浆过程,促进同化物的积累和籽粒灌浆。本研究中,随着施氮量的增加,灌浆速率先升高后降低,在200 kg·hm^-2施氮量下最大灌浆速率和平均灌浆速率达到最大。与单施氮肥处理相比,氮肥和化控共同作用缩短了达到最大灌浆速率时的天数,显著提高了平均灌浆速率和最大灌浆速率,其中,200 kg·hm^-2施氮量下施用化控时灌浆速率最大,达到最大灌浆速率的天数最短。说明化学调控能够在氮肥提高灌浆速率的基础上,进一步改善了籽粒灌浆进程,并在施氮量200 kg·hm^-2和化控剂处理下各籽粒灌浆参数最好,对灌浆的促进作用最明显。

植物内源激素对玉米籽粒灌浆起着重要的调控作用。IAA和CTK可以促进细胞的分裂,灌浆初期IAA和CTK通过促进胚乳细胞分裂,增加了库活性,从而提高玉米籽粒的灌浆速率^[39,40]。籽粒中ABA含量与胚乳细胞增殖速率及籽粒灌浆速率呈极显著正相关,ABA可以促进同化物向籽粒库的运输,促进籽粒灌浆^[41,42]。灌浆前期籽粒较高的GA含量对胚的快速扩增有重要促进作用^[43]。但徐云姬等^[41]研究表明,在胚乳细胞活跃增殖期或籽粒活跃灌浆期,籽粒中GA含量与胚乳细胞的增殖速率或灌浆速率呈极显著负相关。本研究中,灌浆前期和中期籽粒IAA、CTK、GA和ABA含量与灌浆速率显著正相关,说明灌浆前、中期各激素含量的增加有利于提高籽粒灌浆速率。在花后10—30 d,200 kg·hm^-2施氮量下IAA、CTK、GA和ABA含量最高;与单施氮肥处理相比,在氮肥和化学调控共同作用下籽粒各激素含量显著提高,说明化学调控能够在氮肥的促进作用下,进一步提高籽粒各内源激素含量。本研究表明,施氮量200 kg·hm^-2下施用化控剂处理通过提高灌浆前期和中期籽粒IAA、CTK和ABA含量,提高了灌浆速率,从而促进籽粒灌浆。

3.3 高密度下化学调控和氮肥处理对产量及光热水利用效率的影响

前人研究表明,产量增益主要是通过密植群体对光、温、水等资源利用效率的提升最终实现了产量的提高^[17]。本研究相关分析表明,产量与RUE、HUE和WUE呈显著正相关关系,说明较高的光热水利用效率有利于产量的提高。合理的栽培措施,有利于发挥密植增产效应,挖掘玉米生产潜力^[6]。徐昭等^[44]研究表明,适量施氮有利于促进冠层发育,提高光截获量和光合生产力,显著提高RUE和产量。增施氮肥可以促进作物对水分吸收利用,最终提高作物产量^[16]。化控作为作物高产的有效措施,可提高光、热、水利用效率,是实现作物高产的有效途径^[29,45]。王畅等^[29]研究表明,化控显著提高了大豆的RUE和HUE,并提高了产量。本研究中,高密度种植下,随施氮量的增加,玉米RUE、HUE和WUE呈先升高后降低的趋势,产量、穗粒数和粒重均在施氮量200 kg·hm^-2下达到最大,说明200 kg·hm^-2施氮量有利于增加玉米的光热水利用效率,提高产量。与单施氮肥处理相比,氮肥和化学调控共同作用下显著提高了玉米RUE、HUE和WUE,有利于玉米生产中对光热水资源的充分利用,对产量、穗粒数和粒重有明显提高。本研究中,施氮量200 kg·hm^-2下施用化控剂处理有利于实现高密度下玉米资源的高效利用与高产。

4 结论

化控显著提高了叶片P_n和F_v/F_m,增加了籽粒内源激素含量和灌浆速率,提高了光热水利用效率和产量。在200 kg·hm^-2施氮量下,玉米光合特性最好,籽粒ABA、CTK和IAA含量最高,有效增强了灌浆速率,提高了群体光热水利用效率,从而提高了玉米产量。因此,高种植密度下200 kg·hm^-2施氮量配施化控可以改善玉米光合和籽粒灌浆进程,提高光热水利用效率和产量。

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中国农业科学, 2017,50(19):3690-3701.

DOI:10.3864/j.issn.0578-1752.2017.19.006 URL [本文引用: 1]

【Objective】The purpose of this study is to elucidate morphological and physiological mutual beneficial mechanism for compact type maize hybrid under nitrogen and density interaction, for further raise overall productivity of density tolerant maize hybrid.【Method】Plant morphological trait, ear leaf photosynthetic ability and yield were determined under field experimental condition in 2014 and 2015. Split-split plot design, 2 plant type hybrids (compact plant type and flat plant type) as the main plot, 3 nitrogen treatments (N1: 0, N2: 90 kg N·hm^-2 and N3: 180 kg N·hm^-2) as the split plot, 3 plant densities (D1: 45 000 plant/hm², D2: 60 000 plant/hm² and D3: 75 000 plant/hm²) as the sub-split plot. 【Result】The effects of nitrogen on internode length, leaf angle, SPAD value, kernel weight per ear and yield were stronger than that of density on those parameters. Stem diameter, Pn and kernel number per ear was sensitive to density increasing. Compared with flat type hybrid, decreased range of stem diameter was small, and response sensitivity from 1 to 3 internode length was slowness with plant density increased for compact type hybrid. However, the 1-3 internode length was shortened significantly with nitrogen input amount increased (P_N2→N3=0.004-0.028), negative response range of 4-5 internode length for compact type hybrid (10.9%) was higher than positive response range of 4-5 internode length for flat type hybrid (3.3%). Leaf angle of compact type hybrid was down to 2.9°±1.1° with nitrogen input. The leaf angle of leaf below ear leaf changed to a relatively lower with plant density increased. Response peak value of SPAD to nitrogen for compact type hybrid (N₃) was higher than that for flat type hybrid (N₂). The negative effect of Pn caused by density increasing was relatively small for compact type hybrid. SPAD and Pn of ear leaf for compact type hybrid were higher than that for flat type hybrid in N₃ and D₃ treatment. Altogether, the effect of nitrogen and density interaction on kernel number and kernel weight per ear for compact type hybrid was smaller than that for flat type hybrid. Harvest index of compact type hybrid was relatively high, which the difference between N×D interaction treatment (P_N1→N3 =0.16,P_D1→D3 =0.12) was no significant, however, that the difference between that (P_N1→N3 =0.03,P_D1→D3<0.01) of flat type hybrid was very significant. The highest yield record was obtained in N₃D₃ and N₃D₁treatments for compact and flat type hybrid, respectively. And their yield gain ratio for density and nitrogen was 1﹕2.3 and 1﹕4.0, respectively. 【Conclusion】 Compared with flat type hybrid, compact type hybrid had a more adaptable ability of regulating cross and longitudinal growth of basal part of stem. Nitrogen application could reduce leaf angle of leaf above ear leaf, ear leaf and leaf below ear leaf, which could enhance ear leaf light use efficiency. Proper morphophysiological coordinate ability keeps a higher dry matter transfer rate for the compact type hybrid under higher density and higher nitrogen fertilizer condition at kernel weight formation stage, thus achieving a higher population yield.

XIAO W

, LIU

, SHI

, ZHAO H

, WANG Y

. Effect of nitrogen and density interaction on morphological traits, photosynthetic property and yield of maize hybrid of different plant types
Scientia Agricultura Sinica, 2017,50(19):3690-3701. (in Chinese)

DOI:10.3864/j.issn.0578-1752.2017.19.006 URL [本文引用: 1]

李少昆, 赵久然, 董树亭, 赵明, 李潮海, 崔彦宏, 刘永红, 高聚林, 薛吉全, 王立春, 王璞, 陆卫平, 王俊河, 杨祁峰, 王子明. 中国玉米栽培研究进展与展望
中国农业科学, 2017,50(11):1941-1959.

DOI:10.3864/j.issn.0578-1752.2017.11.001 URL [本文引用: 1]

Maize is the first major crop in China and in the world, it plays an important role in ensuring China’s food security. At present, in the face of the rapid development of economic society and a series of problems such as population growth and land reduction, resources shortage and ecological environment deterioration, maize cultivation science is facing new historic opportunities and challenges. In this crucial historical juncture, it is of great significance to review the scientific research and technical progress of maize cultivation in China and to explore the future development direction. Analysis shows that, the aim of maize cultivation research has been transformed from yield production to collaborative development of high yield, high quality, high efficiency, eco-friendly, security and other goals after 60 years of efforts. The research contents were gradually widened and further deepened with remarkable Chinese characteristics. Since entering into the 21th century, the research of maize cultivation has entered a golden development stage. In this stage, a series of breakthroughs in maize cultivation theory, key technology innovation and application have been achieved, which have taken a positive role in ensuring China’s food security. According to the demand of maize production for science and technology in the future and the development trend of modern science and technology, this article indicated that, in the future, high quality, high efficiency, eco-friendly, security will still be the main objectives of maize cultivation. In this article, the key directions and tasks of maize cultivation research in the next 20 years were put forward: (1) Continue to explore the potential of maize yield in different ecological areas and technologies that can realize these potentials, and make every effort to raise the level of yield per unit; (2) Transform the mode of production and take the improving efficiency of resource utilization and labor productivity as goals, reduce the production costs, improve product quality and the market competitiveness of maize; to develop silage and fresh maize so as to promote the diversified development of maize production; (3) In order to respond to the global climate change, carry out the theoretical and technological researches on yield stability and anti-disaster to realize the sustainable production of maize; (4) Based on modern information technology to carry out the researches of intelligent cultivation technology to achieve maize precise production and management; (5) Strengthen the basic researches of maize cultivation and tamp the researches on maize science and technology and the basement of maize production.

LI S

, ZHAO J

, DONG S

, ZHAO

, LI C

, CUI Y

, LIU Y

, GAO J

, XUE J

, WANG L

, WANG

, LU W

, WANG J

, YANG Q

, WANG Z

. Advances and prospects of maize cultivation in China
Scientia Agricultura Sinica, 2017,50(11):1941-1959. (in Chinese)

DOI:10.3864/j.issn.0578-1752.2017.11.001 URL [本文引用: 1]

LIU Y

, HOU

, XIE R

, HAO W

, LI S

, MEI X

. Spatial variation and improving measures of the utilization efficiency of accumulated temperature
Crop Science, 2015,55(4):1806-1817.

DOI:10.2135/cropsci2014.10.0735 URL [本文引用: 1]

[12]

TAO Z

, LI C

, LI J

, DING Z

, XU,

, SUN X

, ZHOU P

, ZHAO

. Tillage and straw mulching impacts on grain yield and water use efficiency of spring maize in Northern Huang-Huai-Hai Valley
Crop Journal, 2015,3(5):445-450.

DOI:10.1016/j.cj.2015.08.001 URL [本文引用: 1]

[13]

BISWAS D

, MA B

. Effect of nitrogen rate and fertilizer nitrogen source on physiology, yield, grain quality, and nitrogen use efficiency in corn
Canadian Journal of Plant Science, 2016,96(3):392-403.

DOI:10.1139/cjps-2015-0186 URL [本文引用: 1]

[14]

ZHAO D

, REDDY K

, KAKANI V

, READ J

, CARTER G

. Corn (Zea mays L.) growth, leaf pigment concentration, photosynthesis and leaf hyperspectral reflectance properties as affected by nitrogen supply
Plant and Soil, 2003,257:205-217.

DOI:10.1023/A:1026233732507 URL [本文引用: 1]

[15]

CHEN J

, YANG Z

, ZHOU

, HAI M

, TANG T

, LIANG Y

, AN T

. Biomass accumulation and partitioning, photosynthesis, and photosynthetic induction in field-grown maize (Zea mays L.) under low-and high-nitrogen conditions
Acta Physiologiae Plantarum, 2013,35(1):95-105.

DOI:10.1007/s11738-012-1051-6 URL [本文引用: 1]

[16]

陈晓璐, 李耕, 刘鹏, 高辉远, 董树亭, 王振林, 张吉旺, 赵斌. 6-苄氨基嘌呤(6-BA)对不同氮素水平下玉米叶片衰老过程中光系统 II性能的调控效应
作物学报, 2013,39(6):1111-1118.

DOI:10.3724/SP.J.1006.2013.01111 URL [本文引用: 2]

A two years experiment was conducted using maize inbred line Qi 319 with different nitrogen fertilization levels and exogenous hormone 6 benzyl adenine (6-BA) treatments. The results demonstrated that the chlorophyll content and net photosynthetic rate (P_n) were gradually decreased after anthesis. In treatments of 6-BA without nitrogen fertilizer, chlorophyll contents were increased significantly, but P_n showed no significant increase, and the activity of PSII reaction center was not increased. Under the condition of applying nitrogen fertilizer, P_n and the activity of PSII reaction center were increased significantly, and based on nitrogen fertilizer used, the 6-BA could improve the response of P_n and performance of PSII to nitrogen fertilizer effectively. Under the nitrogen fertilizer and 6-BA interaction conditions, the chlorophyll content and P_n were increased significantly (P<0.05), the activities of donor site and acceptor site of PSII were increased significantly, and the donor site was improved more than that of acceptor site. The most obvious improvement of the activity of PSII reaction center was appeared at the 10^th day after anthesis. The improvement of photosynthetic performance significantly improved the single plant dry matter accumulation and grain yield (P<0.05). In conclusion, spraying plant with 6-BA under suitable application of nitrogen fertilizer can significantly improve the photosynthetic performance of maize leaf.

CHEN X

, LI

, LIU

, GAO H

, DONG S

, WANG Z

, ZHANG J

, ZHAO

. Effects of exogenous hormone 6 Benzyl Adenine (6-BA) on photosystem II performance of maize during process of leaf senescence under different nitrogen fertilization levels
Acta Agronomica Sinica, 2013,39(6):1111-1118. (in Chinese)

DOI:10.3724/SP.J.1006.2013.01111 URL [本文引用: 2]

朴琳, 任红, 展茗, 曹凑贵, 齐华, 赵明, 李从锋. 栽培措施及其互作对北方春玉米产量及耐密性的调控作用
中国农业科学, 2017,50(11):1982-1994.

DOI:10.3864/j.issn.0578-1752.2017.11.004 URL [本文引用: 2]

【Objective】The purpose of this study was to investigate the regulating effect of cultivation measures and their interactions on grain yield and density resistance of spring maize hybrids, and its contribution to increase of grain yield.【Method】Maize cultivar “Zhongdan 909” was used as experimental materials in 2013 and 2014, which exhibited high yield in the high plant population. From 45 000 plants/hm² to 10 5000 plants/hm², five plant population treatments were designed. Subsoiling (S), wide-narrow planting (W) and chemical regulator (C) as cultivation measures, and composed different cultivation modes by split-split-plot design. Path analysis, factor regression and ANOVA analysis of different cultivation modes based on the yield, and using stepwise regression to analyze the efficiency of resource utilization factors under different cultivation modes, combined with the meteorological data. 【Result】The chemical regulator (C) had a significantly positive effect on yield in the integrated measures mode (contribution rate, 27%-41%), which the effect rests with the plant density increasing by 11 700 plants/hm²under only chemical regulator treatment; wide-narrow planting (W) showed obvious different effects among the treatments. However, the effect of subsoiling (S) on yield displayed priority to indirect effect (contribution rate, 24%-37%), nevertheless, subsoiling plus wide-narrow planting compared with tradition mode (RU) could increase yield by 11.28%. The yield improvement of multiple measures interaction was much higher than those of double measures interaction and a single measure. Compared with traditional mode, multiple measures, double measures and a single measure increased yield by 31.27%, 15.57% and 7.96%, respectively, in a normal year (2013); and increase yield by 15.02%, 11.32% and 5.65%, respectively, in a drought year (2014). The yield increasing was mainly due to the increased population density, and coordinated regulation among radiation use efficiency (RUE), growth degree days use efficiency (GUE) and nitrogen partial factor productivity, then achieved the high yield and high efficiency under integrated measures. 【Conclusion】The yield improvement of multiple measure interaction mode (SWC) was the highest, compared to the traditional mode, the multiple measures could increase plant density by 62 700 plants/hm² and obtain yield improvement by 11.91%, which the improvement was mainly attributed to the optimized population density under multiple measures interaction and regulating effect from integrated measures on resources utilization efficiency of intensive spring maize.

PIAO

, REN

, ZHAN

, CAO C

, QI

, ZHAO

, LI C

. Effect of cultivation measures and their interactions on grain yield and density resistance of spring maize
Scientia Agricultura Sinica, 2017,50(11):1982-1994. (in Chinese)

DOI:10.3864/j.issn.0578-1752.2017.11.004 URL [本文引用: 2]

MOMEN

, KOOCHEK

, MAHALLATI M

. Analysis of the variations in dry matter yield and resource use efficiency of maize under different rates of nitrogen, phosphorous and water supply
Journal of Plant Nutrition, 2020,43(9):1306-1319.

DOI:10.1080/01904167.2020.1729802 URL [本文引用: 1]

[19]

AGGARWAL

, SINGH

, SINGH S

, KANG J

. Heat utilization vis-a-vis crop performance of mechanically transplanted rice (Oryza sativa L.) as affected by tillage systems and nitrogen levels
Journal of Agrometeorology, 2015,17(1):84-89.

[本文引用: 1]

[20]

KITONYO O

, SADRAS V

, ZHOU

, DENTON M

. Nitrogen supply and sink demand modulate the patterns of leaf senescence in maize
Field Crops Research, 2018,225:92-103.

DOI:10.1016/j.fcr.2018.05.015 URL [本文引用: 1]

[21]

WEI S

, WANG X

, SHI D

, LI Y

, ZHANG J

, LIU

, ZHAO

, DONG S

. The mechanisms of low nitrogen induced weakened photosynthesis in summer maize (Zea mays L.) under field conditions
Plant Physiology and Biochemistry, 2016,105:118-128.

DOI:10.1016/j.plaphy.2016.04.007 URLPMID:27101123 [本文引用: 1]

Soil nitrogen (N) shortage is a problem which affects many developing nations. Crops grown with low soil N levels show a marked decrease in the rate of photosynthesis and this deficiency reduces crop yield significantly. Therefore, developing a better understanding of the mechanisms by which low N levels cause decreased photosynthesis is crucial for maize agriculture. To better understand this process, we assessed the responses of photosynthesis traits and enzymatic activities in the summer maize cultivar Denghai 618 under field conditions with and without the use of N fertilisers. We measured photosynthesis parameters, and compared proteome compositions to identify the mechanisms of physiological and biochemical adaptations to N deficiency in maize. We observed that parameters that indicated the rate of photosynthesis decreased significantly under N deficiency, and this response was associated with leaf senescence. Moreover, we identified 37 proteins involved in leaf photosynthesis, and found that N deficiency significantly affected light-dependent and light-independent reactions in maize leaf photosynthesis. Although further analysis is required to fully elucidate the roles of these proteins in the response to N deficiency, our study identified candidate proteins which may be involved in the regulatory mechanisms involved in reduced photosynthesis under low N conditions in maize.

[22]

JIN

, CUI

, LI

, ZHANG J

, DONG S

, LIU

. Effects of integrated agronomic management practices on yield and nitrogen efficiency of summer maize in North China
Field Crops Research, 2012,134:30-35.

DOI:10.1016/j.fcr.2012.04.008 URL [本文引用: 1]

Excessive nitrogen (N) fertilizer application, outdated fertilization techniques, and non-optimal planting patterns are current cultivation problems with summer maize (Zea mays L.) among smallholders in North China. To try to solve those problems, this study examined four integrated agronomic management treatments-MT (FP: traditional farming practices; OPT-1: an optimized combination of cropping systems and fertilizer treatment; HY: treatment based on high-yield studies; and OPT-2: further optimized combination of cropping systems and fertilizer treatment) and four N application rate treatments-NT (0, 129.0, 184.5, and 300.0 kg ha(-1)). Maize grain yield and N efficiency were determined under each treatment. Grain yield, yield components, individual/population dry matter weight, N partial factor productivity (PFPN), N use efficiency (NUE), and agronomic efficiency of N (AEN) were measured. Results from the NT revealed no significant increase in grain yield in response to N applied above 184.5 kg ha(-1) and increased yield was achieved by the MT. In MT. the change in sowing method from relay intercropping to direct seeding was effective in avoiding maize rough dwarf virus and in encouraging mechanized production; individual and population dry matter accumulation from the tasseling stage (VT) to physiological maturity stage (R6) increased in response to delayed sowing date and harvesting time; grain yield increased significantly from more ears per hectare due to increased planting density; and high N efficiency was achieved after optimizing fertilization patterns. In this study, OPT-2 obtained 67.0%, 104.0%, and 53.5% higher grain yield, PFPN, and NUE, respectively, compared to FP, achieving high yield and high N efficiency. Furthermore, the low AEN value suggests that further reduction in the N application rate of OPT-2 may be possible. (C) 2012 Elsevier B.V.

[23]

王庆燕, 管大海, 潘海波, 李建民, 段留生, 张明才, 李召虎. 油菜素内酯对春玉米灌浆期叶片光合功能与产量的调控效应
作物学报, 2015,41(10):1557-1563.

DOI:10.3724/SP.J.1006.2015.01564 URL [本文引用: 1]

A field experiment was conducted to investigate the effect of plant growth regulator brassinolide (BR) on the regulation of physiological activities of ear leaf, the grain filling and yield in spring maize (Zea mays L., cv. Zhengdan 958). Husks and ear leaves were treated with BR at 15 days after anthesis by foliar spraying. The results showed that the chlorophyll content, photosynthetic rate and activities of PEP carboxylaes, RuBP carboxylase, sucrose synthase and sucrose phosphate synthase in ear leaf as well as source content in kernel were significantly reduced, while starch accumulation and dry weight of kernel were increased during grain filling. BR significantly increased the content of chlorophyll and photosynthetic rate, enhanced the activities of PEP carboxylaes and RuBP carboxylase, definitely resulting in enhanced CO₂ assimilation. BR markedly increased the activities of sucrose synthase and sucrose phosphate synthase in ear leaf. And in kernel, the accumulation of sucrose and starch as well as the grain filling rate were accelerated by BR. BR treatment significantly reduced the length of ear barren tip, and increased the number of kernels per ear, which led to higher yield. These results suggest that BR treatment at filling stage may be a good strategy to get a high yield in maize, by enhancing activity of resource, delaying sencence and stimulating the ?ow of assimilate from the source to the sink.

WANG Q

, GUAN D

, PAN H

, LI J

, DUAN L

, ZHANG M

, LI Z

. Effect of brassinolide on leaf photosynthetic function and yield in spring maize filling stage
Acta Agronomica Sinica, 2015,41(10):1557-1563. (in Chinese)

DOI:10.3724/SP.J.1006.2015.01564 URL [本文引用: 1]

边大红, 张瑞栋, 段留生, 李建民, 李召虎. 局部化控夏玉米冠层结构、荧光特性及产量研究
华北农学报, 2011,26(3):139-145.

DOI:10.7668/hbnxb.2011.03.028 URL [本文引用: 1]

Canopy structure has strong effects on chlorophyll fluorescence characteristic and grain yield in maize(Zea mays L.) production.In this paper,the effects of partial spraying of plant growth regulator(PGR),which was sprayed at initial stage of stem elongation at intervals of three lines,were studied on canopy structure,chlorophyll fluorescence parameters and yield characters using a half-compact maize variety,Jinhai 5.The results showed that the plants sprayed PGR had lower plant heights,shorter lengths of internodes under ear,longer lengths of internodes above ear and thicker diameters of basal stems.Therefore,compared to the control plants,a′waving′ canopy was shaped in the treatment sprayed PGR partially.It is found that the ′waving′ canopy could obviously improve light permeating conditions in the middle and lower layers,increase leaf area index and hold longer duration,keep higher chlorophyll content,PS Ⅱ potential activity(Fv/Fo) and chloroplast PS Ⅱ photochemical efficiency,and result in an increase of ear number per area,100-grain weight and yield.

BIAN D

, ZHANG R

, DUAN L

, LI J

, LI Z

. Effects of partial spraying of plant growth regulator on canopy structure, chlorophyll fluorescence characteristic and yield of summer maize ( Zea mays L.)
Acta Agriculturae Boreali-Sinica, 2011,26(3):139-145. (in Chinese)

DOI:10.7668/hbnxb.2011.03.028 URL [本文引用: 1]

王晓慧, 张磊, 刘双利, 曹玉军, 魏雯雯, 刘春光, 王永军, 边少锋, 王立春. 不同熟期春玉米品种的籽粒灌浆特性
中国农业科学, 2014,47(18):3557-3565.

DOI:10.3864/j.issn.0578-1752.2014.18.004 URL [本文引用: 1]

【Objective】 A field experiment was conducted to illustrate grain-filling characteristics of maize hybrids differing in maturities in northeast China, aimed to provide scientific information for the regulation of grain weight and selection of maturity.【Method】 Forty high-yield hybrids differing in maturities were used as experimental materials, and they were planted according to 60 000 plants/hm² with the same environmental conditions. The days of seedling emergence to maturity were investigated, and the whole growing days were applied to divide the maturity of different maize hybrids. The difference of yield and 100-kernel weight of maize hybrids differing in maturities was studied, and the grain-filling processes of maize hybrids differing in maturities were analyzed by Logistic model. The grain-filling parameters of initial grain-filling potential, grain-filling rate and grain-filling duration and their relation to yield were calculated. 【Result】 The results indicated that forty maize hybrids were divided into 4-type maturities according to the whole growing days on the basis of the criterion of Food and Agriculture Organization of the United Nations, and the type I is medium maturity, the type II is mid-late maturity, the type III is late maturity, the type IV is supper-late maturity, respectively. The ratio of type III was 50% and 121-130 d with the whole growing days, the highest among these 4 types, the type I and II were the lowest (12.5%), 101-110 d and 111-120 d with the whole growing days, the ratio of type IV was 25% and 131-140 d with the whole growing days. The yield of late maturity was all the highest among these 4 types, the second of supper-late maturity and the lowest in medium maturity, lower 45%, 44% and 35% than the late maturity, supper-late maturity and mid-late maturity. The yield of mid-late maturity, late maturity and supper-late maturity were not different significantly (P＞0.05), but remarkably higher than medium maturity under trial density (P＜0.05). Initial stage of grain-filling, the 100-kernel weight of medium maturity was the lowest, and highest in the supper-late maturity. After 30 days of grain-filling, the 100-kernel growth rate of mid-late maturity was the fastest among these 4 types, and the slowest in the medium maturity. The tendency of grain-filling rate of these 4 types was expressed in single peak curve; the peak value of medium maturity was the highest, and the lowest in the supper-late maturity. After the peak of grain-filling, the grain-filling rate of supper-late maturity decreased the slowest, and fell the fastest in the medium maturity. The grain-filling processes of maize hybrids differing in maturities were analyzed by Logistic model, and the total filling period could be divided into early stage, middle stage and late stage. The initial grain-filling potential (R₀), maximum grain-filling rate (R_max), grain-filling duration of early stage (T₁), grain-filling rate of middle stage and late stage (v₂, v₃) of medium maturity were significantly higher than other three types in this trial, but the active grain-filling period (P), grain-filling duration of middle stage and late stage (T₂, T₃), grain-filling rate of early stage (v₁) were remarkably lower than the other three types. Furthermore, correlation analysis showed that active grain-filling period, effective grain-filling time (t₃), grain-filling duration of early stage, grain-filling rate of middle stage and late stage were significantly and positively correlated to yield, and initial grain-filling potential, maximum grain-filling rate, grain-filling duration of early stage, grain-filling rate of middle stage and late stage were significantly and negatively correlated to yield. 【Conclusion】 Medium maturity varieties were quickly in launch of grain-filling, short time of active grain-filling period and effective grain-filling time, but the mid-late maturity, late maturity and supper-late maturity varieties were slowly in launch of grain-filling, long time of active grain-filling period and effective grain-filling time. Prolonging the active grain-filling period, effective grain-filling time, grain-filling duration of middle stage and late stage and enhancing the mean grain-filling rate of early stage can improve grain yield of maize hybrids differing in maturities.

WANG X

, ZHANG

, LIU S

, CAO Y

, WEI W

, LIU C

, WANG Y

, BIAN S

, WANG L

. Grain filling characteristics of maize hybrids differing in maturities
Scientia Agricultura Sinica, 2014,47(18):3557-3565. (in Chinese)

DOI:10.3864/j.issn.0578-1752.2014.18.004 URL [本文引用: 1]

陈传永, 王荣焕, 赵久然, 徐田军, 王元东, 刘秀芝, 刘春阁, 裴志超, 成广雷, 陈国平. 不同生育时期遮光对玉米籽粒灌浆特性及产量的影响
作物学报, 2014,40(9):1650-1657.

DOI:10.3724/SP.J.1006.2014.01650 URL [本文引用: 2]

CHEN C

, WANG R

, ZHAO J

, XU T

, WANG Y

, LIU X

, LIU C

, PEI Z

, CHENG G

, CHEN G

. Effects of shading on grain-filling properties and yield of maize at different growth stages
Acta Agronomica Sinica, 2014,40(9):1650-1657. (in Chinese)

DOI:10.3724/SP.J.1006.2014.01650 URL [本文引用: 2]

 

[27]

ZHAO D

, HAN Y

, FU Y

. Changes of plant hormones in inflorescence during sex determination in maize
Acta Phyophsiologica Sinica, 1999,12(2):57-65.

[本文引用: 1]

[28]

GAO

, LIANG X

, ZHANG

, LIN

, ZHAO

, ZHOU L

, SHEN

, ZHOU S

. Spraying exogenous 6-benzyladenine and brassinolide at tasseling increases maize yield by enhancing source and sink capacity
Field Crops Research, 2017,211:1-9.

DOI:10.1016/j.fcr.2017.05.027 URL [本文引用: 1]

[29]

王畅, 赵海东, 冯乃杰, 郑殿峰, 梁晓艳, 齐德强, 黄文婷. 两个生态区大豆光热资源利用率和产量的差异及对化控剂的响应
应用生态学报, 2018,29(11):3615-3624.

DOI:10.13287/j.1001-9332.201811.021 URLPMID:30460808 [本文引用: 3]

The field experiment was conducted at two farms at Jiusan in Heihe (the fourth accumulated temperature zone) and at Lindian County of Daqing (the second accumulated temperature zone), both sites located in Heilongjiang Province, China. With soybean Kenfeng 41 as the test material, uniconazole (S3307, 50 mg.L(-1)) and 2-N, N- diethylamino ethyl caproate (DTA-6, 50 mg.L(-1)) were sprayed on leaves in the early flowering period of soybean. Through grey correlation analysis, the main factors affecting soybean yield were examined, and the differences of the light and heat utilization efficiency and soybean yield in two ecological conditions were compared. The regulation effects of chemical control technology on the light and heat utilization efficiency of soybean were explored. The results showed that the total surface radiation and >/=10 effective accumulated temperature were the main factors affecting soybean yield in both areas compared with rainfall and sunshine hours. The light and heat utilization efficiency from sowing to flowering period was significantly positively correlated with dry matter accumulation, and that from flowering to podding period was significantly positively correlated with dry matter accumulation per plant. There was a significant positive correlation between yield and dry matter accumulation, grain number per plant, grain mass per plant and 100-grain mass at seedling stage to podding stage. S3307 and DTA-6 could significantly improve light and heat utilization efficiency and soybean yield in both areas. S3307 showed the better regulation function to impact the light and heat utilization efficiency and yield than DTA-6 in both sites. In the two ecological areas of Jiusan and Lindian, spraying S3307 increased light utilization efficiency by 13.6% and 17.1%, and increased heat utilization efficiency by 14.1% and 17.2%, respectively. The yield by spraying S3307 was increased by 14.1% and 17.3% separately in Jiusan and Lindian. Therefore, it is the effective way to enhance resources utilization and achieve high-yield by using the reasonable chemical control technology.

WANG

, ZHAO H

, FENG N

, ZHENG D

, LIANG X

, QI D

, HUANG W

. Differences in light and heat utilization efficiency and yield of soybean in two ecological zones and their response to chemical control regulators
Chinese Journal of Applied Ecology, 2018,29(11):3615-3624. (in Chinese)

DOI:10.13287/j.1001-9332.201811.021 URLPMID:30460808 [本文引用: 3]

MIRANZADEH

, EMAM

, SEYYED

, ZARE

. Productivity and radiation use efficiency of four dryland wheat cultivars under different levels of nitrogen and chlormequat chloride
Journal of Agricultural Science and Technology, 2011,13(3):339-351.

[本文引用: 1]

[31]

STICHLIKOVA

, HEJNAK

, SAFRANKOVA

. The effect of abscisic acid and benzylaminopurine on photosynthesis and transpiration rates of maize (Zea mays L.) under water stress and subsequent rehydration
Cereal Research Communications, 2007,35(4):1593-1602.

DOI:10.1556/CRC.35.2007.4.6 URL [本文引用: 1]

[32]

张莉, 赵雪, 曲令华, 王佳慧, 吕新月, 林珊, 周顺利. 外源ABA处理对离体玉米子粒灌浆特性的影响
玉米科学, 2017,25(4):67-74.

[本文引用: 1]

ZHANG

, ZHAO

, QU L

, WANG J

, Lü X

, LIN

, ZHOU S

. Effects of ABA application on maize grain-filling properties in vitro culture
Journal of Maize Sciences, 2017,25(4):67-74. (in Chinese)

[本文引用: 1]

[33]

LONG S

, ZHU X

, NAIDU S

, ORT D

. Can improvement in photosynthesis increase crop yields?
Plant, Cell and Environment, 2006,29(3):315-330.

URLPMID:17080588 [本文引用: 1]

[34]

魏廷邦, 柴强, 王伟民, 王军强. 水氮耦合及种植密度对绿洲灌区玉米光合作用和干物质积累特征的调控效应
中国农业科学, 2019,52(3):428-444.

DOI:10.3864/j.issn.0578-1752.2019.03.004 URL [本文引用: 1]

【Objective】 In oasis irrigation agricultural region, some problems has caused serious influenced of maize production, such as soil available water and nitrogen hunger, premature senescence and unreasonable planting density. To provide technical support for high and stable maize yield, the effects of different ratio of application irrigation and nitrogen and planting density on photosynthesis, dry matter accumulation characteristics and maize yield were studied. 【Method】 Photosynthetic ability, dry matter accumulation characteristics and yield were determined under two-years field experiment, which was carried out in Hexi Oasis irrigation region of Gansu province from 2016 to 2017. In this research, the cultivar “Xianyu335” was applied as research material. A split-split plot design was used as this experiment, with two irrigation application amount treatments (namely 4 050 m ³·hm ^-2 (W₁) and 3 720 m ³·hm ^-2 (W₂)) as the main plot, three nitrogen application amount treatments (namely 0 (N₀), 300 kg·hm ^-2(N₁) and 450 kg·hm ^-2 (N₂)) as the split plot, and three plant densities (namely 7.5×10 ⁴ plant/hm ²(D₁), 9.75×10 ⁴ plant/hm ²(D₂) and 1.2×10 ⁵ plant/hm ²(D₃)) as the split-split plot. 【Result】 Nitrogen fertilizer application and planting density had significant influence on photosynthetic rate, maximum dry matter accumulation rate, emergence days of maximum dry matter accumulation rate, dry matter accumulation amount, grain yield, water use efficiency and nitrogen fertilizer use rate in growth stages of maize. The coupling of irrigation and nitrogen fertilizer management increased photosynthesis, the highest dry matter accumulation rate and advanced the days of emergence of the highest dry matter accumulation rate, and enhanced dry matter accumulation amount and grain yield in growth stages of maize. Under the reduced 20% irrigation and the level of higher nitrogen application in growth stages of maize, compared with the low planting density and high planting density treatments, the photosynthetic rate under the medium planting density treatment was increased by 17.31% and 11.43%, respectively. While, compared with the low planting density treatment, the maximum dry matter accumulation rate and days of emergence of the highest dry matter accumulation rate under the treatment with the high planting density and medium planting density was increased by 21.07% and 7.52%, respectively, and advanced by 6.7, 4.1 days, respectively, meanwhile, the dry matter accumulation of the high planting density treatment was increased by 4.27% and 10.59%, respectively; Compared with the low planting density treatment and the high planting density treatment, the grain yield, water use efficiency and nitrogen fertilizer use rate of maize with the medium planting density treatment was increased by 24.2%, 11.4%, 29.9% and 29.2%, 18.4%, 13.8%, respectively. Under the reduced 20% irrigation and same planting density treatment in growth stages of maize, compared with medium nitrogen application treatment and no nitrogen application treatment, the photosynthetic rate, the dry matter accumulation and grain yield of maize under the treatment with high nitrogen application treatment was increased by 7.34%, 11.63%, 14.63% and 49.54%, 44.53%, 69.03%, under the medium planting density treatment, respectively; Compared with medium nitrogen application treatment and no nitrogen application treatment, the maximum dry matter accumulation rate and days of emergence of the highest dry matter accumulation rate of maize with the high nitrogen application treatment was increased by 19.07% and 54.35% and advanced by 3.9 and 6.8 days under the high planting density treatment, respectively. Compared with no nitrogen application treatment, nitrogen fertilizer use rate of maize with the high nitrogen application treatment was increased by 24.50%. The facts showed that the coupling of reduced 20% irrigation and high nitrogen application had regulated dry matter accumulation, grain yield with the improvement of photosynthesis, dry matter accumulation rate, water use efficiency, nitrogen fertilizer use rate and extending the duration of dry matter accumulation. 【Conclusion】 The treatment with application coupling of irrigation and nitrogen (i.e. reduced 20% irrigation amount during growth 3 720 m ³·hm ^-2(W₂) and N application with 450 kg·hm ^-2 at growth stage and medium density of 9.75×10 ⁴ plant/hm ² at growth stage of maize) could be considered as the best feasible cultivation pattern management, which could provide technical guidance for further exploring high yield and efficient cultivation of close planting maize in Oasis irrigation region.

WEI T

, CHAI

, WANG W

, WANG J

. Effects of coupling of irrigation and nitrogen application as well as planting density on photosynthesis and dry matter accumulation characteristics of maize in oasis irrigated areas
Scientia Agricultura Sinica, 2019,52(3):428-444. (in Chinese)

DOI:10.3864/j.issn.0578-1752.2019.03.004 URL [本文引用: 1]

王红军, 郭书亚, 张艳, 尚赏. 不同密度条件下化控对夏玉米光合特性及产量的影响
江苏农业科学, 2017,45(17):66-69.

[本文引用: 1]

WANG H

, GUO S

, ZHANG

, SHANG

. Effects of chemical control on photosynthetic characteristics and yield of summer maize under different density conditions
Jiangsu Agricultural Sciences, 2017,45(17):66-69. (in Chinese)

[本文引用: 1]

[36]

聂乐兴, 姜兴印, 吴淑华, 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.

URLPMID:21328943 [本文引用: 1]

采用田间小区试验，以高产玉米新品种登海661为材料，研究了拔节期叶面喷施10、20和40 mg·L^-1的胺鲜酯（DA-6）对玉米叶片光合羧化酶、保护酶活性和产量的影响.结果表明：喷施胺鲜酯各处理玉米分别比对照（含有表面活性剂和水）增产10.0%(10 mg·L^-1)、8.9%(20 mg·L^-1)和9.4%(40 mg·L^-1)，增产效果显著，但各浓度间差异不显著.胺鲜酯处理后，花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(P＜0.05)，且对光合速率、RuBP羧化酶和PEP羧化酶活性的影响随着处理浓度的增加而提高；与对照相比，胺鲜酯处理后吐丝期、灌浆期、乳熟期和蜡熟期叶片SOD、CAT、POD和GSTs活性及可溶性蛋白质含量显著提高(P＜0.05)，MDA含量显著降低(P＜0.05)，其中CAT活性随着处理浓度的增加呈上升趋势，其余生理指标各浓度间无显著性差异.

NIE L

, JIANG X

, WU S

, ZHANG J

, 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)

URLPMID:21328943 [本文引用: 1]

赵春玲, 王秀萍, 董朋飞, 刘天学. 化学调控对玉米抗倒防衰及产量和品质的影响
河南农业科学, 2013,42(5):41-44.

URL [本文引用: 1]

为降低连阴雨灾害诱发的玉米倒伏、早衰和产量损失,采用矮壮素浸种以及矮壮素和玉黄金叶面喷施3个处理,研究了植物生长调节物质对玉米叶片衰老、倒伏、籽粒产量和品质的影响。结果表明,3种处理均在一定程度上提高了吐丝期和吐丝后20d玉米叶片的SPAD值、超氧化物歧化酶(SOD)活性,降低了叶片丙二醛(MDA)含量和根茎倒伏率。籽粒中蛋白质、淀粉和脂肪含量无显著变化,赖氨酸含量显著增加。矮壮素浸种和叶面喷施处理分别比清水对照增产2.87%和7.35%,叶面喷施玉黄金对籽粒产量无显著影响。

ZHAO C

, WANG X

, DONG P

, LIU T

. Effects of chemical regulators on lodging, senescence, grain yield and quality of maize under stress environment
Journal of Henan Agricultural Sciences, 2013,42(5):41-44. (in Chinese)

URL [本文引用: 1]

申丽霞, 王璞, 张软斌. 施氮对不同种植密度下夏玉米产量及子粒灌浆的影响
植物营养与肥料学报, 2005,11(3):314-319.

DOI:10.11674/zwyf.2005.0305 URL [本文引用: 1]

Maize grain yield was closely associated with kernel number at maturity, and kernel number was affected by apical kernel abortion mainly occurred at early grain filling stage. The objectives of this work were (i) to evaluate the (effect) of N on grain yield and its components, and to determine the relationship between grain yield and above-ground (dry) matter accumulation; (ii) to analyze the difference of kernel development between apical and mid-basal kernels at grain filling stage, and explore the possible reasons affecting kernel development. The field experiment with summer maize (Zea mays L.) hybrid Zhengdan 958 were carried out at Wuqiao experiment station of China Agricultural University in 2003. The treatments consisted of different crop densities( 52500 plant/ha, 67500 plant/ha, 82500 plant/ha) and nitrogen supply rates (0 kg/ha, 120 kg/ha, 240 kg/ha, 360 kg/ha). Grain yield and its components, above-ground dry matter accumulation, grain filling and content of soluble carbohydrates in summer maize were studied. The results showed that the grain yield per plant was mainly depended on the number of kernels per ear at maturity while the kernel weight was relatively constant. The grain yield and kernel number were closely associated with above-ground dry matter accumulation apparently improved by nitrogen supply. Grain filling and metabolism of soluble carbohydrates in apical and mid-basal kernels were apparently different. In 5—20 day after pollination(DAP), the size of fresh grain, grain dry weight, grain filling rate, total content of soluble sugar, content of sucrose and starch in apical kernels were lower than these in the mid-basal kernels, while in apical kernels these were higher in nitrogen supply treatments than in treatments without nitrogen supply. The grain filling rate in apical kernels began to (decrease) in 15—20 DAP while kernel abortion usually occurred. The results suggested that one of the main reasons affecting development of apical and mid-basal kernels was assimilate supply. Assimilate supply in apical kernels could be improved by nitrogen supply and kernel abortion were inhibited.

SHEN L

, WANG

, ZHANG R

. Effect of nitrogen supply on yield and grain filling in summer maize with different crop density
Plant Nutrition and Fertilizer Science, 2005,11(3):314-319. (in Chinese)

DOI:10.11674/zwyf.2005.0305 URL [本文引用: 1]

SINGH

, GURUNG S

. Hormonal role in the problem of sterility in Oryza sativa
Plant Physiology and Biochemistry, 1982,9:22-32.

[本文引用: 1]

[40]

王纪华, 王树安, 赵冬梅, 梁振兴. 玉米籽粒发育的调控研究Ⅰ. 不同基因型玉米籽粒发育特点及其生理生化基础
中国农业科学, 1996,29(3):33-40.

URL [本文引用: 1]

对不同基因型玉米籽粒发育过程中若干生理生化指标的测定结果表明：果穗顶部大量败育的典型秃尖品种与对照相比，在授粉后4d可见吲哚乙酸(IAA)、玉米素核苷（ZR）达到较高的水平，但在授粉后8～12d却显著低于对照；8～12d出现较高的赤霉酸（GA#-3）和低的脱落酸（ABA）含量。其中授粉后4～8d秃尖品种顶部籽粒的GA#-3/ABA由3.5迅速升高到5.5,而不秃顶部籽粒的该值由3.0下降到1.5左右；授粉后4～28d秃尖顶部籽粒及穗轴中的可溶性酸性蔗糖转化酶活性显著低于对照，以上可能是导致其籽粒败育的重要生理生化原因。

WANG J

, WANG S

, ZHAO D

, LIANG Z

. A study on regulation of maize kernels Ⅰ. characteristics of development of maize kernels from different genotypes and the physiological and biochemical basis
Scientia Agricultura Sinica, 1996,29(3):33-40. (in Chinese)

URL [本文引用: 1]

徐云姬, 顾道健, 张博博, 张耗, 王志琴, 杨建昌. 玉米果穗不同部位籽粒激素含量及其与胚乳发育和籽粒灌浆的关系
作物学报, 2013,39(8):1452-1461.

DOI:10.3724/SP.J.1006.2013.01452 URL [本文引用: 2]

Kernels at the upper position of a maize ear usually show poorer filling and lower weight in contrast to those at the basal position. The mechanism has not been understood. The experiments were conducted using a maize cultivar Denghai 11, grown in both field and greenhouse conditions, to determine number of endosperm cells, kernel filling rate, and contents of zeatin (Z) + zeatin riboside (ZR), indole-3-acetic acid (IAA), abscisic acid (ABA), and gibberellin-3 (GA₃) in the kernels at different positions on an ear from silking to maturity. The results indicated that the maximum number of endosperm cells, maximum division rate of endosperm cells,mean endosperm cell division rate,maximumkernel filling rate, mean kernel filling rate and 100-kernel weight for kernels showed an order of basal position > middle position> upper position. During the active endosperm development and the active kernel filling period, contents of Z+ZR, IAA and ABA in kernels were the greatest at the basal position, the mediate at the middle position, and the least at the upper position on an ear. The two experiments exhibited similar results. The endosperm cell division rate and kernel filling rate were very significantly and positively correlated with contents of Z+ZR, IAA, and ABA in kernels, whereas significantly and negatively correlated with content of GA₃. The results suggested that a lower kernel weight at the upper position on a maize ear is mainly due to a smaller division rate of endosperm cells, leading to less number of endosperm cells, which is closely associated with lower contents of Z+ZR, IAA, and ABA, as well as higher content of GA₃ in the kernels during kernel filling.

XU Y

, GU D

, ZHANG B

, ZHANG

, WANG Z

, YANG J

. Hormone contents in kernels at different positions on an ear and their relationship with endosperm development and kernel filling in maize
Acta Agronomica Sinica, 2013,39(8):1452-1461. (in Chinese)

DOI:10.3724/SP.J.1006.2013.01452 URL [本文引用: 2]

ZEEVAART J A

, CREELMAN R

. Metabolism and physiology of abscisic acid
Annual Review Plant Physiology and Plant Molecular Biology, 1988,39:439-473.

DOI:10.1146/annurev.pp.39.060188.002255 URL [本文引用: 1]

[43]

PHANG T

, SHAO G

, LIAO

, YAN X

, LAM H

. High external phosphate (Pi) increases sodium ion uptake and reduces salt tolerance of ‘Pi-tolerant' soybean
Physiologia Plantarum, 2009,135(4):412-425.

DOI:10.1111/j.1399-3054.2008.01200.x URLPMID:19210751 [本文引用: 1]

Previous studies on the interaction between environmental inorganic phosphate (Pi) and salinity stress using soybean cultivars sensitive to high external Pi had two limitations: (1) the phenotype was dominated by overaccumulation of phosphorus (P); and (2) no detailed analysis was performed for sodium ion uptake. In this study, we focused on the effects of high external Pi on the sodium ion uptake in 'Pi-tolerant' soybean cultivars. The P accumulation in Pi-tolerant soybean Union was much lower [9.0 mg g(-)(1) dry weight (DW); contrasting to 38-76 mg g(-)(1) DW in the 'Pi-sensitive' soybean cultivars]. At in planta level, high level of external Pi significantly (P < 0.001) increased net sodium ion uptake and aggravated salinity stress symptoms. The effects of high external Pi diminished when de-rooted plants were used, suggesting that root is the primary organ interacting with Pi in the growth medium. Two-cell models, including soybean suspension cells and the tobacco Bright Yellow-2 cell line, were also employed to study the effects of high external Pi at the cellular level. Consistent to in planta results, high external Pi uplifted cellular sodium ion uptake and reduced cell viability under salinity stress. Gene expression analyses further showed that HPi (2 mM Pi supplements; excessive level of Pi) could reduce the fold of induction of GmSOS1 and GmCNGC under salinity stress, suggesting that they may be possible molecular targets involved in the interaction between high external Pi and Na(+) uptake.

[44]

徐昭, 史海滨, 李仙岳, 田彤, 付小军, 李正中. 水氮限量供给对盐渍化农田玉米光能利用与产量的影响
农业机械学报, 2018,49(12):281-291.

[本文引用: 1]

, SHI H

, LI X

, TIAN

, FU X

, LI Z

. Effect of limited irrigation and nitrogen rate on radiation utilization efficiency and yield of maize in salinization farmland
Transactions of the Chinese Society for Agricultural Machinery, 2018,49(12):281-291. (in Chinese)

[本文引用: 1]

[45]

彭亚静, 汪新颖, 张丽娟, 郝晓然, 乔继杰, 王玮, 吉艳芝. 根层调控对小麦-玉米种植体系氮素利用及土壤硝态氮残留的影响
中国农业科学, 2015,48(11):2187-2198.

DOI:10.3864/j.issn.0578-1752.2015.11.010 URL [本文引用: 1]

【Objective】 Low resource utilization efficiency of water and fertilizer and high soil nitrogen accumulation were common problems of the winter wheat-summer maize system in the North China Plain. This study was conducted to investigate the effects of different root layer regulation measures on nitrogen utilization and soil NO₃^--N residue of crops. 【Method】 The field experiment was conducted in the high-yield farmlands in North China Plain. Traditional water-nitric nitrogen, water-nitric nitrogen regulation, regulation+Agh (soil conditioner), regulation+CRU (controlled-release urea), regulation+GGR (plant growth regulator) were designed as five treatments. Soil, plant and root samples were collected and measured, and the nitrogen utilization effects of different root layer regulation measures were analyzed.【Result】The results indicated that under the premise of controlling water and nitrogen, the wheat-maize yield and nitrogen uptake in Agh and GGR treatments were higher than traditional water and nitrogen treatments. At jointing stage of wheat, roots at 80-100 cm soil layers in GGR treatment distributed much, which shows that GGR could promote the development of roots at lower layer. At bell stage of maize, at 20-50 cm layer, root length density of Agh treatment in Gaocheng and GGR treatment in Daming was significantly higher than those of traditional water and nitrogen treatment. In the first rotation season, in 0-200 cm layer, the soil residual nitrate-N in GGR treatment in Gaocheng and Shenzhou was significantly lower than that of traditional water and nitrogen treatment, especially at 60-100 cm layer soil residue nitrate nitrogen was the lowest. In the second rotation season, the soil residual nitrate-N in Agh treatment in Gaocheng and GGR treatment in Daiming were significantly lower than traditional water and nitrogen treatment. The amount of apparent deficiency of nitrogen of Agh in the first rotation season and GGR (Gaocheng) in the second rotation season was large, which indicate that root regulation could promote crop to absorb soil accumulated nitrogen. Root layer regulation measures were of benefits to economy and ecology, Water Use Efficiency (WUE) and Partial Factor Productivity_Nitrogen (PFP_N) increased by 2.47 kg·m^-3 and 18.08 kg·kg^-1 compared with traditional water and nitrogen treatment, and the average income was 258.43 yuan/667 m². 【Conclusion】 In high yield land in the North China Plain, different root-layer regulation measures of single cropping and annual production of wheat and maize respectively increased by 8.58%, 5.99% and 7.13% in average compared with the traditional water and nitrogen treatment. After two seasons the residue of soil nitrate nitrogen in 0-100 cm layer soil were reduced by 70.73 kg·hm^-2 and 59.44 kg·hm^-2 in average compared with the conventional water and nitrogen treatment, obviously reduced the residue of soil nitrate nitrogen, slowed the leaching loss of nitrogen to deep soil, and promoted root development during critical growing period of wheat-maize. All in all, Agh and GGR were effective measures to improve crops yield, use the soil accumulated nitrogen, achieve cost synergies and improve water and fertilizer use efficiency.

PENG Y

, WANG X

, ZHANG L

, HAO X

, QIAO J

, WANG

, JI Y

. Effects of root layer regulation on nitrogen utilization and soil NO₃^--N residue of wheat-maize system
Scientia Agricultura Sinica, 2015,48(11):2187-2198. (in Chinese)

DOI:10.3864/j.issn.0578-1752.2015.11.010 URL [本文引用: 1]