化肥减量条件下秸秆还田配施腐植酸对玉米碳氮代谢的影响

郭伟; 但武侠; 马传芳; 孙海燕

doi:10.19303/j.issn.1008-0384.2023.04.012

化肥减量条件下秸秆还田配施腐植酸对玉米碳氮代谢的影响

doi: 10.19303/j.issn.1008-0384.2023.04.012

郭伟^{1, 2,},
但武侠¹,
马传芳¹,
孙海燕^{1, 3}

1.
黑龙江八一农垦大学农学院，黑龙江　大庆　163319
2.
黑龙江省现代农业栽培技术与作物种质改良重点实验室，黑龙江　大庆　163319
3.
农业农村部东北平原农业绿色低碳重点实验室，黑龙江　大庆　163319

基金项目: 黑龙江省应用技术研究与开发计划项目（GA20B102-02）；黑龙江八一农垦大学基础培育课题（ZRCPY202101）

详细信息

作者简介:
郭伟（1977−），男，教授，博士生导师，主要从事作物生理生态研究（E-mail：agrigw@163.com）

中图分类号: S513
计量
- 文章访问数: 429
- HTML全文浏览量: 246
- PDF下载量: 19
- 被引次数: 0
出版历程
- 收稿日期: 2022-08-21
- 修回日期: 2023-03-03
- 网络出版日期: 2023-05-09
- 刊出日期: 2023-04-28

Effects of Chemical Fertilizer Reduction with Humic Acid Addition on Carbon and Nitrogen Metabolism of Zea mays under Straw-returning Cultivation

GUO Wei^{1, 2
,},
DAN Wuxia¹,
MA Chuanfang¹,
SUN Haiyan^{1, 3}

1.
College of Agriculture, Heilongjiang Bayi Agricultural University, Daqing, Heilongjiang　163319, China
2.
Heilongjiang Provincial Key Laboratory of Modern Agricultural Cultivation and Germplasm Improvement, Daqing, Heilongjiang　163319, China
3.
Key Laboratory of Low Carbon Green Agriculture in Northeast Plain，Ministry of Agriculture and Rural Affairs, Daqing, Heilongjiang　163319, China

摘要

摘要: 目的秸秆直接全量还田技术在东北黑土区农业生产中被广泛应用，但关于还田秸秆腐解过程对当季作物的直接影响及其配套技术研究较少。对秸秆与腐植酸互作效应的研究可为完善东北黑土区秸秆还田技术提供参考。方法在减少15%化肥常规施用量的基础上，以化肥常量施肥（CF）为对照，设置化肥减量15%增施腐植酸（HA）、化肥减量15%增施秸秆（SR）、化肥减量15%增施腐植酸和秸秆（SRH）处理，研究玉米秸秆还田配施腐植酸对当季玉米植株碳氮代谢的影响，以期明确秸秆还田及腐植酸对化肥减量情况下玉米生长的生理效应。结果秸秆还田提高了玉米叶片磷酸烯醇式丙酮酸羧化酶（PEPC）活性，腐植酸提高了玉米叶片1,5-二磷酸核酮糖羧化酶（Rubisco）和PEPC活性，秸秆还田配施腐植酸在PEPC活性上具有互作效应。虽然秸秆还田降低了蔗糖磷酸合成酶（SPS）活性，但腐植酸提高了SPS活性，两者均提高了蔗糖合成酶活性，且两者互作下同步提高了酸性转化酶（AI）和中性转化酶（NI）活性，促进了还原糖（RS）合成，进而提高了总可溶性糖（TSS）含量，加强了淀粉合成。秸秆还田对玉米植株氮代谢无直接影响，腐植酸对氮代谢关键酶活性具有正效应。结论秸秆还田配施腐植酸提高了化肥减量条件下玉米叶片光合效率和光合产物的贮存能力，植株氮代谢水平不低于常规施肥对照。
- 秸秆还田 /
- 化肥减量 /
- 腐植酸 /
- 碳氮代谢
Abstract: Objective Effects on plant metabolism of the widely practiced utilizing spent straws for fertilization in northeastern China corn fields were studied. Method In a field experimentation, application of either the conventional chemical fertilization (CK) or 15% reduction of chemical fertilizer with an addition of humic acid (HA), spent straws (SR), or both HA and SR (SRH) was implemented on a black soil lot of Zea mays. Physiological responses including carbon and nitrogen metabolisms to the treatments of current-season corn plants were monitored. Result The replacement of chemical fertilizer by SR enhanced the phosphoenolpyruvate carboxylase (PEPC) activity, that by HA decreased the 1,5-ribulose bisphosphate carboxylase (Rubisco) and PEPC activities, and that by SRH interacted on PEPC in the corn leaves. Although SR decreased, HA increased the sucrose phosphate synthase (SPS) activity. Both SR and HA (SRH) treatment raised the activities of sucrose synthase (SS), and combined use of HA and HA(SRH) increased the acid invertase (AI), and neutral invertase (NI) promoting the synthesis of reducing sugars that resulted in an increased total soluble sugar (TSS) and starch synthesis. SR had no significant effect, but HA had a positive effect on key enzymes involving the nitrogen metabolism of the plant. Conclusion With a 15% reduction on chemical fertilizer, addition of SRH boosted the photosynthetic efficiency and storage of photosynthetic products of corn leaves without any significant effect on the nitrogen metabolism of the plant.
- Straw-returning /
- chemical fertilizer reduction /
- humic acid /
- carbon and nitrogen metabolisms

HTML全文

图 1 化肥减量时秸秆还田配施腐植酸对碳氮比的影响

不同小写字母表示同一时期不同处理之间差异显著（P＜0.05）。图2同。

Figure 1. Effects of straw-returning with HA addition on carbon/nitrogen ratio of plants under chemical fertilizer reduction

Data with different letters indicate significant differences between treatments in same period (P＜0.05). Same for Fig.2.

下载: 全尺寸图片幻灯片

图 2 化肥减量时秸秆还田配施腐植酸对植株干重的影响

Figure 2. Effects of straw-returning with HA addition on dry mass of plant under chemical fertilizer reduction

下载: 全尺寸图片幻灯片

表 1 试验处理的各物料用量及代号

Table 1. Amounts of materials applied and treatment codes

试验处理 Experimental treatments	化肥 Chemical fertilizer/(kg·hm⁻²)			秸秆 Straw/ (kg·hm⁻²)	腐植酸 Humic acid/ (kg·hm⁻²)
试验处理 Experimental treatments	尿素 Urea	磷酸二铵 Diammonium phosphate	硫酸钾 Potassium sulphate	秸秆 Straw/ (kg·hm⁻²)	腐植酸 Humic acid/ (kg·hm⁻²)
CF	375	225	150	0	0
HA	320	190	0	0	600
SR	320	190	0	3750	0
SRH	320	190	0	3750	600

下载: 导出CSV

表 2 化肥减量时秸秆还田配施腐植酸对玉米碳代谢的影响

Table 2. Effect of straw-returning cultivation with HA addition on carbon metabolism of corn under chemical fertilizer reduction

指标 Indexes	处理 Treatments	2019年		2020年
指标 Indexes	处理 Treatments	拔节期 JS	抽雄期 TS	拔节期 JS	抽雄期 TS
1,5二磷酸核酮糖羧化酶活性 Rubisco activity/( U·g⁻¹)	CF	182.478±19.869 c	168.553±11.127 b	107.010±7.919 c	82.530±9.306 c
	HA	333.133±31.305 a	246.361±23.953 a	160.524±13.897 a	214.903±19.634 a
	SR	187.207±12.636 c	178.220±20.805 b	97.900±10.600 c	146.339±15.249 b
	SRH	295.363±28.578 b	250.515±21.954 a	128.584±10.431 b	209.210±21.617 a
磷酸烯醇式丙酮酸羧化酶活性 PEPC activity/( U·g⁻¹)	CF	179.762±23.198 c	186.567±21.519 d	124.569±10.988 c	238.509±8.438 c
	HA	231.241±30.347 b	263.827±19.643 c	150.838±14.381 c	285.123±36.233 b
	SR	267.463±26.890 b	321.121±42.789 b	175.993±16.885 b	315.329±31.267 b
	SRH	340.904±30.898 a	360.571±18.653 a	202.089±20.991 a	432.205±43.422 a
蔗糖磷酸合成酶活性 SPS activity/( U·g⁻¹)	CF	492.938±48.295 c	479.661±60.974 b	458.702±70.678 b	492.480±75.921 a
	HA	996.230±94.786 a	633.180±78.493 a	734.351±48.045 a	577.201±24.592 a
	SR	529.382±53.381 bc	444.858±27.624 b	467.885±53.758 b	315.727±36.193 b
	SRH	622.420±76.004 b	550.510±57.402 ab	482.118±50.025 b	505.940±68.491 a
蔗糖合成酶活性 SS activity/( U·g⁻¹)	CF	0.558±0.046 c	0.435±0.017 b	0.384±0.043 c	0.333±0.041 c
	HA	0.643±0.054 bc	0.439±0.065 b	0.481±0.044 b	0.404±0.055 b
	SR	0.692±0.072 b	0.493±0.036 b	0.481±0.044 b	0.462±0.023 ab
	SRH	0.951±0.086 a	0.718±0.105 a	0.577±0.063 a	0.532±0.013 a
酸性转化酶活性 AI activity/( U·g⁻¹)	CF	0.188±0.011 c	0.260±0.025 c	0.605±0.030 a	0.415±0.047 b
	HA	0.257±0.023 b	0.317±0.029 b	0.708±0.070 a	0.484±0.034 a
	SR	0.242±0.025 b	0.364±0.050 b	0.635±0.068 a	0.497±0.071 a
	SRH	0.310±0.038 a	0.426±0.049 a	0.680±0.058 a	0.539±0.059 a
中性转化酶活性 NI activity/( U·g⁻¹)	CF	0.119±0.016 b	0.201±0.011 c	0.394±0.036 b	0.383±0.039 b
	HA	0.146±0.016 a	0.231±0.028 c	0.484±0.037 a	0.433±0.056 ab
	SR	0.145±0.016 a	0.266±0.030 b	0.487±0.046 a	0.421±0.017 ab
	SRH	0.150±0.011 a	0.329±0.022 a	0.517±0.058 a	0.483±0.055 a
还原糖含量 RS content/(mg·g⁻¹)	CF	6.448±0.497 b	10.439±1.021 c	12.707±0.858 b	14.162±2.297 a
	HA	6.715±0.288 ab	12.571±1.730 b	12.707±1.184 b	15.307±1.125 a
	SR	6.795±0.345 ab	13.909±0.923 b	14.271±1.479 ab	14.716±1.295 a
	SRH	7.290±0.600 a	18.290±2.265 a	15.981±0.755 a	17.118±2.281 a
可溶性糖含量 SS content/(mg·g⁻¹)	CF	76.805±2.100 b	183.208±19.136 c	147.315±12.109 b	218.915±13.809 b
	HA	80.668±3.948 b	243.913±25.718 b	177.415±18.147 a	249.490±22.366 ab
	SR	89.401±5.496 a	302.668±18.060 a	169.865±14.699 a	202.365±25.940 b
	SRH	90.295±6.562 a	292.733±31.149 a	177.115±17.130 a	270.490±27.253 a
淀粉含量 Starch content/(mg·g⁻¹)	CF	4.706±0.289 d	5.351±0.503 d	2.002±0.212 c	4.594±0.405 d
	HA	5.890±0.595 c	7.170±0.650 c	4.316±0.339 b	5.449±0.409 c
	SR	11.014±1.217 a	11.692±0.560 a	6.707±0.643 a	8.996±0.773 a
	SRH	8.863±0.884 b	9.385±0.561 b	4.905±0.440 b	7.984±0.629 b
不同字母表示同一时期不同处理间差异显著(P＜0.05)。表4同。 Data with different letters indicate significant differences between treatments in same period (P＜0.05). Same for Table 4.

下载: 导出CSV

表 3 化肥减量时秸秆还田配施腐植酸对玉米碳代谢影响的相关分析

Table 3. Correlation between straw-returning with HA addition and carbon metabolism of corn under chemical fertilizer reduction

指标 Indices	1.5-二磷酸核酮糖羧化酶 Rubisco	磷酸烯醇式丙酮酸 PEPC	蔗糖磷酸合成酶 SPS	酸性转化酶 AI	中性转化酶 NI	蔗糖合成酶 SS	还原糖 RS	总可溶性糖 SS	淀粉 Starch
秸秆还田 SR	−0.002	0.555^**	−0.379^**	0.173	0.180	0.489^**	0.277^*	0.183	0.716^**
腐植酸 HA	0.618^**	0.336^**	0.569^**	0.194	0.158	0.361^**	0.201	0.161	−0.026
1.5-二磷酸核酮糖羧化酶 Rubisco	1.000	0.445^**	0.615^**	−0.495^**	−0.541^**	0.596^**	−0.330^**	−0.126	0.324^**
磷酸烯醇式丙酮酸羧化酶 PEPC	0.445^**	1.000	−0.098	−0.178	−0.037	0.342^**	0.287^*	0.448^**	0.668^**
蔗糖磷酸合成酶 SPS	0.615^**	−0.098	1.000	−0.164	−0.286^*	0.302^*	−0.383^**	−0.320^*	−0.133
酸性转化酶 AI	−0.495^**	−0.178	−0.164	1.000	0.929^**	−0.302^*	0.639^**	0.289^*	−0.329^**
中性转化酶 NI	−0.541^**	−0.037	−0.286^*	0.929^**	1.000	−0.431^**	0.771^**	0.462^**	−0.275^*
蔗糖合成酶 SS	0.596^**	0.342^**	0.302^*	−0.302^*	−0.431^**	1.000	−0.357^**	−0.371^**	0.457^**
还原糖 RS	−0.330^**	0.287^*	−0.383^**	0.639^**	0.771^**	−0.357^**	1.000	0.804^**	0.035
总可溶性糖 SS	−0.126	0.448^**	−0.320^*	0.289^*	0.462^**	−0.371^**	0.804^**	1.000	0.263^*
淀粉 Starch	0.324^**	0.668^**	−0.133	−0.329^**	−0.275^*	0.457^**	0.035	0.263^*	1.000
为P＜0.05水平显著性相关，为P＜0.01水平显著性相关。表5同。 indicates significant difference at P＜0.05; ** significant difference at P＜0.01. Same for Table 5.

下载: 导出CSV

表 4 化肥减量时秸秆还田配施腐植酸对玉米氮代谢的影响

Table 4. Effect of straw-returning cultivation with HA addition on nitrogen metabolism of corn under chemical fertilizer reduction

指标 Indices	处理 Treatments	2019年		2020年
指标 Indices	处理 Treatments	拔节期 JS	抽雄期 TS	拔节期 JS	抽雄期 TS
硝酸还原酶活性 NR activity/(U·g⁻¹)	CF	10.733±0.298 c	15.112±0.604 ab	10.889±0.353 c	22.785±1.491 c
	HA	14.883±0.662 a	15.613±1.300 ab	12.833±0.999 b	25.806±1.561 b
	SR	12.158±0.572 b	14.286±0.961 b	13.381±1.200 b	23.354±1.760 c
	SRH	14.495±1.076 a	16.549±1.244 a	15.911±1.078 a	29.009±2.233 a
亚硝酸还原酶活性 NiR activity/( U·g⁻¹)	CF	7.223±0.397 a	7.855±0.602 ab	5.304±0.427 b	5.617±0.425 ab
	HA	7.467±0.530 a	8.278±0.555 a	5.294±0.534 b	5.781±0.300 ab
	SR	6.066±0.489 b	7.178±0.608 bc	5.209±0.388 b	5.352±0.325 b
	SRH	7.291±0.338 a	8.278±0.647 a	5.937±0.415 a	6.148±0.204 a
谷氨酰胺合成酶活性 GS activity/(U·g⁻¹)	CF	4.335±0.296 c	6.066±0.676 ab	5.063±0.343 a	5.223±0.439 ab
	HA	5.014±0.409 ab	6.946±0.674 a	4.974±0.131 a	5.449±0.405 ab
	SR	4.503±0.497 bc	5.353±0.626 bc	4.846±0.423 a	4.600±0.251 c
	SRH	5.153±0.404 a	6.762±0.785 a	5.342±0.454 a	5.665±0.290 a
谷氨酸合成酶活性 GOGAT activity/(U·g⁻¹)	CF	201.226±12.436 a	168.848±14.888 b	169.267±9.328 a	223.634±15.222 b
	HA	209.045±16.757 a	179.706±12.784 b	177.401±16.887 a	226.755±18.983 b
	SR	157.349±13.598 b	160.202±9.295 b	176.813±13.815 a	180.598±14.901 c
	SRH	201.724±13.005 a	230.400±19.836 a	184.970±18.398 a	291.460±14.460 a
谷氨酸脱氢酶活性 GDH activity/(U·g⁻¹)	CF	154.233±14.485 cd	282.472±25.283 b	122.881±10.086 b	194.298±14.478 c
	HA	189.421±17.192 ab	327.190±32.580 a	151.293±12.562 a	230.185±15.743 b
	SR	174.206±13.139 bc	256.136±19.632 b	138.068±7.485 a	197.765±13.886 c
	SRH	210.899±15.115 a	320.303±12.314 a	146.717±11.322 a	289.820±28.757 a
可溶性蛋白质含量 SP content/(mg·g⁻¹)	CF	2.737±0.133 b	5.389±0.166 b	3.533±0.214 a	6.693±0.553 a
	HA	2.966±0.208 b	5.404±0.242 b	3.690±0.166 a	6.636±0.306 a
	SR	2.980±0.395 b	5.539±0.220 b	3.585±0.216 a	6.029±0.331 b
	SRH	3.490±0.250 a	6.082±0.503 a	3.654±0.201 a	7.020±0.319 a

下载: 导出CSV

表 5 化肥减量时秸秆还田配施腐植酸对玉米氮代谢影响的相关分析

Table 5. Correlation between straw-returning with HA and nitrogen metabolism of corn under chemical fertilizer reduction

指标 Indices	硝酸还原酶 NR	亚硝酸还原酶 NiR	谷氨酰胺合成酶 GS	谷氨酸合成酶 GOGAT	谷氨酸脱氢酶 GDH	可溶性蛋白质 SP
秸秆还田 SR	0.121	−0.056	−0.065	0.048	0.078	0.055
腐植酸 HA	0.259^*	0.266^*	0.404^**	0.450^**	0.328^**	0.103
硝酸还原酶 NR	1.000	−0.172	0.155	0.653^**	0.379^**	0.804^**
亚硝酸还原酶 NiR	−0.172	1.000	0.467^**	0.038	0.642^**	0.059
谷氨酰胺合成酶 GS	0.155	0.467^**	1.000	0.160	0.709^**	0.446^**
谷氨酸合成酶 GOGAT	0.653^**	0.038	0.160	1.000	0.326^**	0.465^**
谷氨酸脱氢酶 GDH	0.379^**	0.642^**	0.709^**	0.326^**	1.000	0.637^**
可溶性蛋白质 SP	0.804^**	0.059	0.446^**	0.465^**	0.637^**	1.000

下载: 导出CSV

参考文献(29)

[1]	XIN L J. Chemical fertilizer rate, use efficiency and reduction of cereal crops in China, 1998–2018 [J]. Journal of Geographical Sciences, 2022, 32(1): 65−78. doi: 10.1007/s11442-022-1936-2
[2]	王囡囡, 朱凤莉, 张春峰, 等. 不同秸秆还田方式对白浆土土壤养分及大豆产量的影响 [J]. 中国土壤与肥料, 2016(1):94−97. doi: 10.11838/sfsc.20160116 WANG N N, ZHU F L, ZHANG C F, et al. Effect of different returning method of straw to soil on soil chemical property and soybean yield [J]. Soil and Fertilizer Sciences in China, 2016(1): 94−97.(in Chinese) doi: 10.11838/sfsc.20160116
[3]	王宇先, 孙士明, 徐莹莹, 等. 黑龙江省半干旱地区生长季玉米秸秆腐解特征研究 [J]. 黑龙江农业科学, 2020(9):46−48. doi: 10.11942/j.issn1002-2767.2020.09.0046 WANG Y X, SUN S M, XU Y Y, et al. Study on decomposition characteristics of maize straw in semi-arid region of Heilongjiang Province during growing season [J]. Heilongjiang Agricultural Sciences, 2020(9): 46−48.(in Chinese) doi: 10.11942/j.issn1002-2767.2020.09.0046
[4]	张水勤, 袁亮, 林治安, 等. 腐植酸促进植物生长的机理研究进展[J]. 植物营养与肥料学报, 2017, 23(4): 1065-1076. ZHANG S Q, YUAN L, LIN Z A, et al. Advances in humic acid for promoting plant growth and its mechanism[J]. Plant Nutrition and Fertilizer Science, 2017, 23(4)4: 1065-1076. (in Chinese)
[5]	SHAHBAZ M, KUMAR A, KUZYAKOV Y, et al. Priming effects induced by glucose and decaying plant residues on SOM decomposition: A three-source ¹³C/¹⁴C partitioning study [J]. Soil Biology and Biochemistry, 2018, 121: 138−146. doi: 10.1016/j.soilbio.2018.03.004
[6]	韦安培, 丁文超, 胡恒宇, 等. 耕作方式及秸秆还田对土壤性质、微生物碳源代谢及小麦产量的影响 [J]. 干旱地区农业研究, 2019, 37(6):145−152. doi: 10.7606/j.issn.1000-7601.2019.06.21 WEI A P, DING W C, HU H Y, et al. Effects of tillage methods and straw return on soil properties, metabolism of microbial carbon source and wheat yield [J]. Agricultural Research in the Arid Areas, 2019, 37(6): 145−152.(in Chinese) doi: 10.7606/j.issn.1000-7601.2019.06.21
[7]	孔雨, 刘格彤, 张颖, 等. 秸秆还田对麦田碳氮代谢生理特性的影响 [J]. 焦作大学学报, 2013, 27(3):82−83. doi: 10.3969/j.issn.1008-7257.2013.03.028 KONG Y, LIU G T, ZHANG Y, et al. Effects of straw returning on physiological characteristics of carbon and nitrogen metabolism in wheat field [J]. Journal of Jiaozuo University, 2013, 27(3): 82−83.(in Chinese) doi: 10.3969/j.issn.1008-7257.2013.03.028
[8]	王娟, 张丽君, 姚槐应. 添加秸秆和黑炭对水稻土碳氮转化及土壤微生物代谢图谱的影响 [J]. 中国水稻科学, 2013, 27(1):97−104. doi: 10.3969/j.issn.1001-7216.2013.01.014 WANG J, ZHANG L J, YAO H Y. Effects of straw and black carbon addition on C-N transformation and microbial metabolism profile in paddy soil [J]. Chinese Journal of Rice Science, 2013, 27(1): 97−104.(in Chinese) doi: 10.3969/j.issn.1001-7216.2013.01.014
[9]	宋蒙亚, 李忠佩, 刘明, 等. 不同有机物料组合对土壤养分和生化性状的影响 [J]. 中国农业科学, 2013, 46(17):3594−3603. doi: 10.3864/j.issn.0578-1752.2013.17.008 SONG M Y, LI Z P, LIU M, et al. Effects of mixtures of different organic materials on soil nutrient content and soil biochemical characteristics [J]. Scientia Agricultura Sinica, 2013, 46(17): 3594−3603.(in Chinese) doi: 10.3864/j.issn.0578-1752.2013.17.008
[10]	李雪利, 叶协锋, 顾建国, 等. 土壤C/N比对烤烟碳氮代谢关键酶活性和烟叶品质影响的研究 [J]. 中国烟草学报, 2011, 17(3):32−36. doi: 10.3969/j.issn.1004-5708.2011.03.007 LI X L, YE X F, GU J G, et al. Effect of soil C/N ratio on activity of key enzymes involved in carbon and nitrogen metabolism and quality of flue-cured tobacco leaves [J]. Acta Tabacaria Sinica, 2011, 17(3): 32−36.(in Chinese) doi: 10.3969/j.issn.1004-5708.2011.03.007
[11]	CRUZ J L, MOSQUIM P R, PELACANI C R, et al. Carbon partitioning and assimilation as affected by nitrogen deficiency in cassava [J]. Photosynthetica, 2003, 41(2): 201−207. doi: 10.1023/B:PHOT.0000011952.77340.53
[12]	高俊凤. 植物生理学实验指导[M]. 北京: 高等教育出版社, 2006.
[13]	任娇, 于寒, 吴春胜, 等. 不同秸秆还田方式下玉米叶片光合、荧光特性及水分利用率的比较 [J]. 分子植物育种, 2017, 15(12):5241−5247. doi: 10.13271/j.mpb.015.005241 REN J, YU H, WU C S, et al. Comparison of photosynthesis, fluorescence characteristics and water use efficiency of maize leaves under different straw returning methods [J]. Molecular Plant Breeding, 2017, 15(12): 5241−5247.(in Chinese) doi: 10.13271/j.mpb.015.005241
[14]	李英浩, 刘景辉, 赵宝平, 等. 干旱胁迫下腐植酸对燕麦叶片非结构性碳水化合物代谢的影响 [J]. 西北植物学报, 2020, 40(12):2101−2107. doi: 10.7606/j.issn.1000-4025.2020.12.2101 LI Y H, LIU J H, ZHAO B P, et al. Effects of humic acid on metabolism of non-structural carbohydrates in oat leaves under drought stress [J]. Acta Botanica Boreali-Occidentalia Sinica, 2020, 40(12): 2101−2107.(in Chinese) doi: 10.7606/j.issn.1000-4025.2020.12.2101
[15]	BATLLE-AGUILAR J, BROVELLI A, PORPORATO A, et al. Modelling soil carbon and nitrogen cycles during land use change. A review [J]. Agronomy for Sustainable Development, 2011, 31(2): 251−274. doi: 10.1051/agro/2010007
[16]	庄振东, 李絮花. 腐植酸氮肥对玉米产量、氮肥利用及氮肥损失的影响 [J]. 植物营养与肥料学报, 2016, 22(5):1232−1239. ZHUANG Z D, LI X H. Effects of humic acid nitrogen fertilization on corn yield, nitrogen utilization and nitrogen loss [J]. Journal of Plant Nutrition and Fertilizers, 2016, 22(5): 1232−1239.(in Chinese)
[17]	谷端银, 王秀峰, 高俊杰, 等. 纯化腐植酸对氮胁迫下黄瓜幼苗生长和氮代谢的影响 [J]. 应用生态学报, 2018, 29(8):2575−2582. doi: 10.13287/j.1001-9332.201808.029 GU D Y, WANG X F, GAO J J, et al. Effects of purified humic acid on the growth and nitrogen metabolism of cucumber seedlings under nitrogen stress [J]. Chinese Journal of Applied Ecology, 2018, 29(8): 2575−2582.(in Chinese) doi: 10.13287/j.1001-9332.201808.029
[18]	LIU J, ZHONG Y, JIA X Y, et al. Wheat straw decomposition patterns and control factors under nitrogen fertilization [J]. Journal of Soil Science and Plant Nutrition, 2021, 21(4): 3110−3121. doi: 10.1007/s42729-021-00592-z
[19]	NARDI S, PIZZEGHELLO D, GESSA C, et al. A low molecular weight humic fraction on nitrate uptake and protein synthesis in maize seedlings [J]. Soil Biology and Biochemistry, 2000, 32(3): 415−419. doi: 10.1016/S0038-0717(99)00168-6
[20]	裴丽珍, 陈远学, 张雯雯, 等. 有机物料还田对夏玉米穗位叶光合性能及氮代谢的影响 [J]. 作物学报, 2022, 48(8):2115−2124. PEI L Z, CHEN Y X, ZHANG W W, et al. Effects of organic material returned on photosynthetic performance and nitro-Gen metabolism of ear leaf in summer maize [J]. Acta Agronomica Sinica, 2022, 48(8): 2115−2124.(in Chinese)
[21]	袁天佑, 王俊忠, 冀建华, 等. 施用腐植酸对夏玉米产量、氮素吸收及氮肥利用率的影响 [J]. 核农学报, 2017, 31(4):794−802. doi: 10.11869/j.issn.100-8551.2017.04.0794 YUAN T Y, WANG J Z, JI J H, et al. Effects of application humic acid on yield, nitrogen absorption and nitrogen use efficiency of summer maize [J]. Journal of Nuclear Agricultural Sciences, 2017, 31(4): 794−802.(in Chinese) doi: 10.11869/j.issn.100-8551.2017.04.0794
[22]	朱祖祥. 从绿肥的起爆效应探讨它的肥效机制及其在施用上的若干问题 [J]. 浙江农业科学, 1963, 4(3):104−109. doi: 10.16178/j.issn.0528-9017.1963.03.002 ZHU Z X. Discussing the fertilizer effect mechanism of green manure and its application problems [J]. Journal of Zhejiang Agricultural Sciences, 1963, 4(3): 104−109.(in Chinese) doi: 10.16178/j.issn.0528-9017.1963.03.002
[23]	WILD B, LI J, PIHLBLAD J, et al. Decoupling of priming and microbial N mining during a short-term soil incubation [J]. Soil Biology and Biochemistry, 2019, 129: 71−79. doi: 10.1016/j.soilbio.2018.11.014
[24]	MAZZILLI S R, KEMANIAN A R, ERNST O R, et al. Priming of soil organic carbon decomposition induced by corn compared to soybean crops [J]. Soil Biology and Biochemistry, 2014, 75: 273−281. doi: 10.1016/j.soilbio.2014.04.005
[25]	MOORHEAD D L, SINSABAUGH R L. A theoretical model of litter decay and microbial interaction [J]. Ecological Monographs, 2006, 76(2): 151−174. doi: 10.1890/0012-9615(2006)076[0151:ATMOLD]2.0.CO;2
[26]	DONG L H, CÓRDOVA-KREYLOS A L, YANG J S, et al. Humic acids buffer the effects of urea on soil ammonia oxidizers and potential nitrification [J]. Soil Biology and Biochemistry, 2009, 41(8): 1612−1621. doi: 10.1016/j.soilbio.2009.04.023
[27]	REEZA A A, AHMED O H, MAJID N M N A, et al. Reducing ammonia loss from urea by mixing with humic and fulvic acids isolated from coal [J]. American Journal of Environmental Sciences, 2009, 5(3): 420−426. doi: 10.3844/ajessp.2009.420.426
[28]	GARCÍA A C, BERBARA R L L, FARÍAS L P, et al. Humic acids of vermicompost as an ecological pathway to increase resistance of rice seedlings to water stress [J]. African Journal of Biotechnology, 2012, 11(13): 3125−3134.
[29]	CANELLAS L P, PICCOLO A, DOBBSS L B, et al. Chemical composition and bioactivity properties of size-fractions separated from a vermicompost humic acid [J]. Chemosphere, 2010, 78(4): 457−466. doi: 10.1016/j.chemosphere.2009.10.018