不同春季追氮模式对小麦茎秆抗倒性能及木质素积累的影响

doi:10.3864/j.issn.0578-1752.2020.21.009

摘要/Abstract

摘要：

【目的】探讨春季不同追氮模式对小麦各节间茎秆抗倒伏能力、木质素积累及籽粒产量的影响,明确高施氮量条件下适宜的春季追氮模式,为小麦高产稳产抗逆应变栽培提供技术支撑。【方法】于2017—2018和2018—2019年2个小麦生长季,以倒伏敏感型品种山农16和抗倒伏品种济麦22为供试材料,在高施氮量300 kg·hm^-2基施1/3条件下设置4种春季追肥模式,分别为等量二次性追氮和剩余一次性追氮,即起身期﹕孕穗期1/3﹕1/3（T1）,拔节期﹕开花期1/3﹕1/3（T2）,孕穗期一次性追施剩余2/3氮（T3）和拔节期一次性追施剩余2/3氮（CK）。深入研究春季不同追氮模式对冬小麦植株茎秆抗折力、木质素积累、木质素合成关键酶基因的表达丰度及籽粒产量的调控效应。【结果】抗倒伏品种济麦22的各节间茎秆抗折力、木质素积累量以及单体含量均高于倒伏敏感型品种山农16,2种类型品种开花期T1、CK处理的抗折力高于T2和T3处理,木质素积累量、单体的含量表现为T1>T3>CK>T2,灌浆期和成熟期各处理间抗折力、木质素积累量以及单体的含量表现为T1>T3>T2>CK。灌浆期山农16和济麦22在T1处理下抗折力较CK、T2、T3处理分别增加24.69%、19.97%、13.15%和26.92%、15.36%、5.87%;山农16和济麦22在T1处理下的各生育阶段木质素积累量平均值分别较CK、T2、T3处理提高了21.71%、15.45%、8.85%和25.19%、21.75%、15.83%;成熟期2个品种T1处理的木质素S型单体含量平均值分别较CK、T2、T3处理高18.82%、18.48%、8.39%。不同追氮模式处理的木质素合成相关酶基因（苯丙氨酸解氨酶:PAL、咖啡酸3氧甲基转移酶:COMT、香豆酸-3-羟基氧化酶:C3H、肉桂酰辅酶 A 还原酶:CCR、肉桂酸4羟化酶:C4H等）表达均随生育进程呈下降趋势,其表达量高低依次为T1>T3>T2>CK。孕穗期追氮处理模式的千粒重高于其他处理,因T1处理可提高穗粒数以及群体有效穗数,其最终籽粒产量较高。同一时期相同处理各节间茎秆木质素积累量、单体含量均呈现为I1>I2>I3>I4>I5的趋势。【结论】高施氮量300 kg·hm^-2基施1/3条件下起身与孕穗期等量二次性追氮模式较其他春季追氮模式处理显著提高了小麦开花后茎秆各节间抗折能力、木质素积累量、木质素合成途径相关酶基因的表达以及籽粒产量。因此,起身与孕穗期等量二次性追氮模式可作为黄淮海麦区高施氮量300 kg·hm^-2基施1/3条件下的春季适宜追氮模式。

关键词: 追氮模式, 抗倒性能, 木质素积累, 籽粒产量, 冬小麦

Abstract:

【Objective】The purpose of this experiment was to explore the effects of different spring nitrogen topdressing modes on stem lodging resistance, lignin accumulation, grain yield of winter wheat, and to identify the appropriate spring nitrogen topdressing modes under the condition of high nitrogen application, so as to provide technical support for high and stable yield and stress-resistant cultivation of winter wheat.【Method】In the two wheat growing seasons of 2017-2018 and 2018-2019, the lodging sensitive variety Shannong 16 and the lodging resistant variety Jimai 22 were used as test materials, and the application rate was 1/3 under high nitrogen application rate of 300 kg·hm^-2. There were four types of spring topdressing modes, which were equal amount of secondary nitrogen topdressing and remaining one-time nitrogen topdressing, namely the rising stage﹕booting stage 1/3﹕1/3 (T1), jointing stage: flowering stage 1/3﹕1/3 (T2), the remaining 2/3 nitrogen was applied at the booting stage (T3) and the remaining 2/3 nitrogen was applied at the jointing stage (CK). The effects of different nitrogen topdressing modes on stem resistance, lignin accumulation, expressive abundance of the key genes involving in lignin biosynthesis pathway and grain yield of winter wheat were studied.【Result】The total lignin accumulation and lignin monomers content of the lodging resistance wheat were both higher than those of the lodging sensitive wheat. The breaking strength under T1, CK was higher than that under T2 and T3, the lignin accumulation and monomer content were T1>T3>CK>T2 in two types cultivar, and the breaking strength, lignin accumulation, monomer content under all treatments at grain filling stage and maturity stage were T1>T3>T2>CK in two types cultivar. The breaking strength of Shannong 16 and Jimai 22 under T1 treatment were increased by 24.69%, 19.97%, 13.15% and 26.92%, 15.36%, 5.87%, respectively, compared with CK, T2, T3 at grain filling stage. The average lignin accumulation of Shannong 16 and Jimai22 under T1 at each growth stage was 21.71%, 15.45% , 8.85% and 25.19%, 21.75%, 15.83% higher than CK, T2, and T3, respectively. The average content of S monomer was 18.82%, 18.48%, and 8.39% higher than CK, T2 and T3 at maturity stage, respectively. The expressive abundance of key genes involved in lignin biosynthesis pathway (phenylalanine ammonia-lyase: PAL, caffeic acid3-o-methytransferase: COMT, coumarate-3-hydroxyl oxidase: C3H, innamoyl Co A reductase: CCR, cinnamate 4-hydroxylase: C4H etc.) decreased with the growth process, that tended to T1>T3>T2>CK under different stage of growth. The 1000-grain weight of the nitrogen topdressing remaining one-time at booting stage was higher than other treatments. T1 treatment could increase the spike number, grain number and yield. The lignin accumulation and monomer content of stem in different internodes during the same stage were I1>I2>I3>I4>I5.【Conclusion】Under the condition of high nitrogen application rate of 300 kg·hm^-2 and basal application rate of 1/3, the same amount of secondary nitrogen topdressing modes treatment at the rising stage and booting stages significantly improved the breaking strength, lignin accumulation, lignin monomer content, the expressive abundance of key genes involved in lignin biosynthesis pathway and yield after anthesis stage, compared with other spring nitrogen topdressing modes. Therefore, the same amount of secondary nitrogen topdressing mode at rising stage and booting stage could be used as an appropriate spring nitrogen topdressing mode under the condition of high nitrogen application rate of 300 kg·hm^-2 and basal application rate of 1/3 in Huang-Huai-Hai plain.

Key words: nitrogen topdressing modes, lodging resistance, lignin accumulation, grain yield, winter wheat

董荷荷, 骆永丽, 李文倩, 王元元, 张秋霞, 陈金, 金敏, 李勇, 王振林. 不同春季追氮模式对小麦茎秆抗倒性能及木质素积累的影响[J]. 中国农业科学, 2020, 53(21): 4399-4414.

DONG HeHe, LUO YongLi, LI WenQian, WANG YuanYuan, ZHANG QiuXia, CHEN Jin, JIN Min, LI Yong, WANG ZhenLin. Effects of Different Spring Nitrogen Topdressing Modes on Lodging Resistance and Lignin Accumulation of Winter Wheat[J]. Scientia Agricultura Sinica, 2020, 53(21): 4399-4414.

图/表 8

表1

表2

不同春季追氮模式对不同抗倒伏型小麦品种各节间抗折力的影响"

年份 Year	节间 Internode	处理 Treatment	SN16			JM22
年份 Year	节间 Internode	处理 Treatment	开花期 Anthesis stage	灌浆期 Filling stage	成熟期 Maturity stage	开花期 Anthesis stage	灌浆期 Filling stage	成熟期 Maturity stage
2017-2018	I1	CK	5.99ab	6.65c	3.82bcd	7.54a	7.79cd	4.36ef
		T1	5.92bc	7.75a	4.76a	7.52a	9.38a	5.38a
		T2	4.85ef	7.03bc	4.00bc	6.83b	8.05cd	4.74de
		T3	5.24de	7.28ab	4.83a	5.89d	8.62b	4.83bcd
	I2	CK	6.47a	5.59ef	3.64cd	7.17ab	6.39fg	4.20fg
		T1	5.62bcd	6.97bc	4.80a	7.03b	8.11c	5.26ab
		T2	4.97e	5.81de	3.95bc	6.29c	7.03e	4.77de
		T3	5.48cd	6.16d	4.06b	5.78de	7.66d	4.78cd
	I3	CK	5.19de	4.61g	3.15fg	5.47efg	5.83hi	4.04fgh
		T1	4.42fg	5.72def	4.12b	5.75de	6.60f	4.61de
		T2	4.40fg	4.68g	3.18fg	5.27fg	6.23fgh	3.82hij
		T3	3.76ij	5.49ef	3.50def	4.22j	6.49f	5.22abc
	I4	CK	4.08ghi	4.02ij	2.70hij	5.54def	4.55k	3.26kl
		T1	4.35fgh	5.31f	3.55de	5.45efg	5.98ghi	4.82bcd
		T2	3.89hi	4.52gh	2.93ghi	5.07gh	5.63ij	3.97gh
		T3	3.35jk	5.30f	3.20efg	4.71hi	5.33j	3.92ghi
	I5	CK	3.20kl	3.46k	1.84k	4.33ij	3.48l	2.68m
		T1	3.40jk	4.47ghi	2.97gh	4.68hi	4.60k	3.53jk
		T2	2.54m	3.74jk	2.47j	4.08jk	4.38k	3.04lm
		T3	2.73lm	4.09hij	2.59ij	3.61k	4.35k	3.56ijk
2018-2019	I1	CK	6.46a	6.93cd	3.92cde	8.07a	7.70bc	4.25de
		T1	6.48a	7.94a	4.99a	7.80a	8.54a	5.31a
		T2	5.36bc	7.31bc	4.28bc	6.85b	8.07ab	4.89b
		T3	5.76b	7.78ab	4.49ab	6.64bc	8.20ab	4.84b
	I2	CK	5.05cd	5.62ef	3.36fgh	5.58e	6.65d	4.13de
		T1	5.45bc	6.67d	4.74ab	5.69e	8.16ab	4.75bc
		T2	4.80de	5.87e	3.93cde	5.04f	7.39c	3.86e
		T3	4.29efghi	6.97cd	3.97cde	4.40gh	7.33c	4.33d
	I3	CK	4.44efg	4.75gh	2.91hi	5.62e	5.45gh	3.88e
		T1	4.64def	5.55ef	4.28bcd	5.55e	6.52de	4.76bc
		T2	3.79ijk	5.15fg	3.60efg	4.48gh	6.07ef	3.94de
		T3	4.11ghi	5.20fg	3.68ef	5.06f	6.12ef	4.36cd
	I4	CK	4.06ghi	4.04ij	2.28j	5.82de	4.65ij	3.30gh
		T1	4.35efgh	5.68ef	3.77def	6.22cd	6.19def	3.82ef
		T2	3.55jk	4.35hi	3.15hi	5.66e	5.45gh	3.33fgh
		T3	3.51k	5.45ef	2.93gh	4.75fg	5.89fg	3.86e
	I5	CK	3.92hij	4.04ij	1.24k	4.63fg	4.07k	3.07gh
		T1	4.22fghi	4.57h	3.13gh	4.71fg	5.13hi	3.33gh
		T2	3.40k	3.62j	3.05h	3.59i	4.45jk	2.95h
		T3	2.58l	4.49hi	2.39ij	4.09hi	5.05hi	3.41fg

表2

图1

图2

表3

图3

表4

表5

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名称	序列-F (5′-3′)	序列-R (5′-3′)
ACTIN	GGGACCTCACGGATAATCTAATG	CGTAAGCGAGCTTCTCCTTTAT
PAL	CATCTTGGAGGGAAGCTCATAC	GACTTGGTGGCAAATCGAATAAC
C4H	GCCGAGAGCAAGATCCTCGT	CGTGCTTCTCCTCCTCCAGG
HCT	TGTCAGCATTGCTCCGTGGA	CCAGGAGCCATGAACACCGG
C3H	CCCATCCTCGGCATCACCAT	CGCCCTTCTCAGTCGTGTCA
CCR	CGTGATGGTGCTGAAGAAAC	CGATCATCGAAGCCGATACA
CAD	GAGGTCGTCAAGATGGAC	CTAGCTCTTTCTCCCTCTG
4CL	TGCACACTGGAGACATTGGC	TTCGAGTTCCGCAGGAGGTA
F5H	AGCTCCCCTCTCTCAAGTGC	GACACAGTCCTCGGCGTTCT
COMT	GATCCATGACAACGAGTCTACC	CGAATCAATCGACGACACAAAC
CSE	GCCAGCAAGGACAAGACCATCA	GCTGAGCCAGGCGAGGATGT

年份 Year	生育时期 Growth stage	木质素积累量 Lignin accumulation	木质素单体含量 Lignin monomer content
年份 Year	生育时期 Growth stage	木质素积累量 Lignin accumulation	S型	S型+G型
2017-2018	开花期Anthesis stage	0.82**	0.61**	0.68**
	灌浆期Filling stage	0.93**	0.63**	0.61**
	成熟期Maturity stage	0.90**	0.90**	0.84**
2018-2019	开花期Anthesis stage	0.75**	0.74**	0.71**
	灌浆期Filling stage	0.91**	0.82**	0.70**
	成熟期Maturity stage	0.85**	0.70**	0.66**

年份 Year	品种 Cultivar	处理 Treatment	穗数 Spike number (×10⁴·hm^-2)	穗粒数 Grain number per spikes	千粒重 1000-grain weight (g)	籽粒产量 Grain yield (kg·hm^-2)
2017-2018	SN16	CK	662ab	37.11ab	40.89b	7833.4c
		TI	688a	38.46a	42.08ab	8387.1a
		T2	652ab	35.47b	41.26b	8066.3b
		T3	628b	38.07ab	44.89a	7997.3bc
	JM22	CK	691a	36.10a	41.78c	8336.8bc
		TI	710a	37.79a	45.02ab	9136.4a
		T2	678a	35.47a	43.82bc	8602.8b
		T3	659a	35.88a	45.67a	8107.8c
2018-2019	SN16	CK	719b	39.35a	44.30b	8035.0c
		TI	756a	39.60a	45.39a	8667.5a
		T2	690bc	38.56a	44.60b	8273.9b
		T3	671c	39.40a	45.55a	8168.4bc
	JM22	CK	747b	42.45ab	45.91b	8630.6bc
		TI	775a	42.73a	46.92a	9523.7a
		T2	716c	37.79c	46.06b	8904.9b
		T3	694c	40.26b	46.86a	8466.8c

	生育时期 Growth stage	抗折力 Breaking strength	木质素积累量 Lignin accumulation	木质素单体含量Lignin monomer content
	生育时期 Growth stage	抗折力 Breaking strength	木质素积累量 Lignin accumulation	S型	G型	S型+G型
籽粒产量 Yield	开花期Anthesis stage	0.66	0.86**	0.71*	0.69*	0.75*
	灌浆期Filling stage	0.82**	0.92**	0.85**	0.82**	0.83**
	成熟期Maturity stage	0.69**	0.91**	0.81**	0.75*	0.77*