氮水添加对黑沙蒿种子功能性状的影响

doi:10.17521/cjpe.2024.0047

植物生态学报 ›› 2024, Vol. 48 ›› Issue (12): 1637-1649.DOI: 10.17521/cjpe.2024.0047 cstr: 32100.14.cjpe.2024.0047

所属专题：植物功能性状

氮水添加对黑沙蒿种子功能性状的影响

马斌¹, 佘维维¹^,³, 秦欢¹, 宣瑞智¹, 宋春阳¹, 袁新月¹, 苗春¹, 刘靓¹, 冯薇¹^,³, 秦树高¹^,⁴, 张宇清¹^,²^,³^,⁵^,^*()

¹北京林业大学水土保持学院, 宁夏盐池毛乌素沙地生态系统国家定位观测研究站, 北京 100083
²北京林业大学林木资源高效生产全国重点实验室, 北京 100083
³北京林业大学林业生态工程教育部工程中心, 北京 100083
⁴北京林业大学水土保持国家林业和草原局重点实验室, 北京 100083
⁵北京林业大学林木育种与生态修复国家工程研究中心, 北京 100083

收稿日期:2024-02-18 接受日期:2024-06-01 出版日期:2024-12-20 发布日期:2024-12-20
通讯作者: *张宇清(zhangyq@bjfu.edu.cn)
基金资助:
国家自然科学基金(U22A20504);国家自然科学基金(32071844);国家自然科学基金(32001373)

Effects of nitrogen and water addition on seed functional traits of Artemisia ordosica

MA Bin¹, SHE Wei-Wei¹^,³, QIN Huan¹, XUAN Rui-Zhi¹, SONG Chun-Yang¹, YUAN Xin-Yue¹, MIAO Chun¹, LIU Liang¹, FENG Wei¹^,³, QIN Shu-Gao¹^,⁴, ZHANG Yu-Qing¹^,²^,³^,⁵^,^*()

¹Yanchi Research Station of the Mu Us Desert, School of Soil and Water Conservation, Beijing Forestry University, Beijing 100083, China
²State Key Laboratory of Efficient Production of Forest Resource, Beijing Forestry University, Beijing 100083, China
³Engineering Research Center of Forestry Ecological Engineering, Ministry of Education, Beijing Forestry University, Beijing 100083, China
⁴Key Laboratory of National Forestry and Grassland Administration on Soil and Water Conservation, Beijing Forestry University, Beijing 100083, China
⁵National Engineering Research Center for Tree Breeding and Ecological Restoration, Beijing Forestry University, Beijing 100083, China

Received:2024-02-18 Accepted:2024-06-01 Online:2024-12-20 Published:2024-12-20
Contact: *ZHANG Yu-Qing(zhangyq@bjfu.edu.cn)
Supported by:
National Natural Science Foundation of China(U22A20504);National Natural Science Foundation of China(32071844);National Natural Science Foundation of China(32001373)

摘要/Abstract

摘要： 种子性状对植物种群繁殖更新具有重要影响, 了解黑沙蒿(Artemisia ordosica)种子性状对氮沉降及降水变化的响应, 可为预测未来气候变化背景下荒漠植物群落演替方向提供理论基础。该研究以宁夏毛乌素沙地连续8年(2015-2022)的野外氮、水控制实验(自然降雨、增水20%、增水40%和0、20、60 kg N·hm^-2·a^-1全因子交互)平台为依托, 测定了氮、水添加对黑沙蒿种子形态、生理和化学性状的影响; 采用培养皿萌发实验并通过观测萌发种子数量计算各萌发指标, 探究了各处理种子在两种培养温度下(25 ℃/15 ℃、20 ℃/10 ℃)的萌发特性。结果表明: (1)氮、水添加及其交互作用均显著改变了黑沙蒿种子质量、长度、宽度、曲率、体积、形状系数等形态性状, 增水使种子变小, 氮添加使种子变大, 增水20%削弱了氮添加对种子变大的作用, 增水40%时, 氮添加使种子变小; (2)常温(25 ℃/15 ℃)下种子萌发率总体高于低温(20 ℃/10 ℃), 低温下, 种子萌发性状受亲本氮、水添加影响更大, 且仅增水20%、40%处理下, 种子萌发率更高; (3)氮、水添加处理下, 种子长、宽、质量等形态大小性状调节扩散和幼苗建立功能, 种子形状系数、2 h吸水率、萌发率和萌发速率共同调节种子萌发时间, 单独增水或氮添加使黑沙蒿种子只倾向于单一功能(扩散建立或萌发)的变异, 而同时增水与氮添加使黑沙蒿种子两类功能均发生变异。研究结果表明, 黑沙蒿种群繁殖更新策略在全球气候变化中有较大的可塑性, 可通过种子性状变异改变在氮沉降及降雨增多情形下的种子扩散建立与萌发功能策略。

关键词: 氮沉降, 降雨增多, 黑沙蒿, 种子性状, 萌发, 繁殖

Abstract:

Aims Seed traits are of great significance for clarifying plant population reproduction and regeneration strategies. Exploring responses of Artemisia ordosica seed traits to nitrogen and water addition can improve our understanding of desert plant community succession in the context of global change.

Methods The study was conducted based on a 8-year field water and nitrogen control experiment (2015-2022) in Mu Us Desert. A full factor interaction experiment of ambient precipitation, 20% water increase, 40% water increase and 0, 20, 60 kg N·hm⁻²·a⁻¹ were carried out to determine the morphological, physiological, chemical and germination traits of A. ordosica seed. Seed germination traits under each treatment at two temperatures (25 °C/15 °C and 20 °C /10 °C) were explored by using the petri dish germination experiment and calculating the germination indexes by observing the number of seedling emergence.

Important findings (1) Nitrogen and water addition, as well as their interactions, significantly influenced A. ordosica seed morphological traits, including seed mass, length, width, curvature, volume, form coefficient and other traits. Water addition resulted in smaller seeds while nitrogen alone led to larger seeds. However, 20% water increase mitigated the effect of nitrogen increase on seed size. Conversely, with a 40% water increase, nitrogen addition caused smaller seeds. (2) Germination percentage was higher at normal temperature (25 °C /15 °C) than that at low temperature conditions (20 °C /10 °C). At low temperature, seed germination traits were more affected by water and nitrogen addition experienced by parental organisms, and when only adding 20% and 40% water, A. ordosica had higher germination percentage. (3) Under the treatments of water and nitrogen addition, the morphological traits such as seed length, width, quality jointly regulated function of dispersal and establishment while form coefficient, 2 h water absorption, germination percentage and germination speed jointly regulated ecological function of germination timing. Adding water or nitrogen alone tent to the variation of only one function (seed dispersal and seedling establishment or seed germination) of A. ordosica seeds, while simultaneous addition of water and nitrogen led to the variation of both functions of A. ordosica seeds. These findings demonstrate that populations of A. ordosica possessed a great adaptability in response to global climate change by altering their functional strategies of seed dispersal, seedling establishment and seed germination.

Key words: nitrogen deposition, increased precipitation, Artemisia ordosica, seed traits, germination, reproduction

马斌, 佘维维, 秦欢, 宣瑞智, 宋春阳, 袁新月, 苗春, 刘靓, 冯薇, 秦树高, 张宇清. 氮水添加对黑沙蒿种子功能性状的影响. 植物生态学报, 2024, 48(12): 1637-1649. DOI: 10.17521/cjpe.2024.0047

MA Bin, SHE Wei-Wei, QIN Huan, XUAN Rui-Zhi, SONG Chun-Yang, YUAN Xin-Yue, MIAO Chun, LIU Liang, FENG Wei, QIN Shu-Gao, ZHANG Yu-Qing. Effects of nitrogen and water addition on seed functional traits of Artemisia ordosica. Chinese Journal of Plant Ecology, 2024, 48(12): 1637-1649. DOI: 10.17521/cjpe.2024.0047

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2024.0047

https://www.plant-ecology.com/CN/Y2024/V48/I12/1637

图/表 8

表1 氮、水添加对黑沙蒿种子性状影响的双因素方差分析结果(F值)

Table 1 Effects of nitrogen and water addition on seed traits of Artemisia ordosica by two-factor ANOVAs (F value)

处理 Treatment	种子形态性状 Seed morphological trait
处理 Treatment	质量 Mass	长度 Length		宽度 Width		曲率 Curvature	体积 Volume	表面积 Surface area		宽长比 Width:length		形状系数 FC
氮添加 Nitrogen addition (N)	19.11^***	1.60^*		18.34^***		0.51	25.50^***	21.60^***		24.90^***		6.24^**
水添加 Water addition (W)	36.63^***	190.41^***		148.70^***		2.26^*	150.20^***	313.50^***		9.82^**		10.71^***
N × W	5.41^**	36.19^***		18.36^***		3.23^*	16.10^***	21.87^***		60.60^***		30.09^***
处理 Treatment	种子生理、化学性状 Seed physiological and chemical traits
处理 Treatment	0.5 h吸水率 0.5 ab (%)		1 h吸水率 1 ab (%)		2 h吸水率 2 ab (%)		碳含量 C content (%)		氮含量 N content (%)		碳氮比 C:N
氮添加 Nitrogen addition (N)	3.58^**		1.23		2.70^*		37.81^***		25.22^***		15.07^***
水添加 Water addition (W)	2.80^*		1.96		7.73^***		19.57^***		8.52^***		13.94^***
N × W	0.69		3.99^**		5.09^***		94.30^***		6.57^***		50.53^***

图1 不同氮、水添加下黑沙蒿种子形态性状变化。N0, 不添加氮; N1, 氮添加20 kg·hm-2·a-1; N2, 氮添加60 kg·hm-2·a-1。W0, 不增水; W1, 增水20%; W2, 增水40%。

Fig. 1 Changes of seed morphological traits of Artemisia ordosica under different nitrogen and water additions. N0, no nitrogen addition; N1, nitrogen addition of 20 kg·hm-2·a-1; N2, nitrogen addition of 60 kg·hm-2·a-1. W0, no water addition; W1, 20% water addition; W2, 40% water addition. 0.5 ab, 0.5-hour water absorption of seeds; 1 ab, 1-hour water absorption of seeds; 2 ab, 2-hour water absorption of seeds; C, seed carbon content; Cur, seed curvature; FC, seed form coefficient; L, seed length; m, seed mass; N, seed nitrogen content; SA, seed surface area; V, seed volume; W, seed width.

表2 黑沙蒿种子萌发性状在两种温度萌发条件下对氮、水添加响应的双因素方差分析(F值)

Table 2 Two-factor analyses of variance (F value) of seed germination traits of Artemisia ordosica in response to nitrogen and water addition at two germination temperature treatments

萌发指标 Germination index	25 ℃/15 ℃			20 ℃/10 ℃
萌发指标 Germination index	氮添加 Nitrogen addition (N)	水添加 Water addition (W)	N × W	氮添加 Nitrogen addition (N)	水添加 Water addition (W)	N × W
幼苗高度 Seedling height	7.23^**	4.82^*	3.26^*	2.11	1.01	10.16^***
萌发指数 Germination index	2.16	1.31	5.02^**	27.50^***	39.11^***	16.67^***
活力指数 Vigor index	6.36^**	0.16	6.24^**	14.42^***	26.38^***	7.60^***
萌发速率 Germination speed	2.46	0.29	2.20	20.46^***	37.67^***	17.15^***
萌发率 Germination percentage	3.77^*	13.46^***	9.83^***	32.81^***	17.12^***	4.39
萌发势 Germination energy	0.42	0.49	2.41	42.42^***	40.88^***	34.42^***

图2 两种温度下黑沙蒿种子日累计萌发数量(平均值±标准误)。A, 20 ℃/10 ℃。B, 25 ℃/15 ℃。N0, 不添加氮; N1, 氮添加20 kg·hm-2·a-1; N2, 氮添加60 kg·hm-2·a-1。W0, 不增水; W1, 增水20%; W2, 增水40%。

Fig. 2 Daily cumulative number of Artemisia ordosica seedling emergence under two germination temperature treatments (mean ± SE). A, 20 °C/10 °C. B, 25 °C /15 °C. N0, no nitrogen addition; N1, nitrogen addition of 20 kg·hm-2·a-1; N2, nitrogen addition of 60 kg·hm-2·a-1. W0, no water addition; W1, 20% water addition; W2, 40% water addition.

图3 氮、水添加对黑沙蒿种子萌发率(A、B)和萌发势(C、D)的影响(平均值±标准误)。A、C, 萌发温度为25 ℃/15 ℃。B、D, 萌发温度为20 ℃/10 ℃。N0, 不添加氮; N1, 氮添加20 kg·hm-2·a-1; N2, 氮添加60 kg·hm-2·a-1。W0, 不增水; W1, 增水20%; W2, 增水40%。

Fig. 3 Effects of water and nitrogen addition on seed germination percentage (GP) (A, B) and germination energy (GE) (C, D) of Artemisia ordosica (mean ± SE). A, C, Germination temperature of 25 °C /15 °C. B, D, Germination temperature of 20 °C /10 °C. N0, no nitrogen addition; N1, nitrogen addition of 20 kg·hm-2·a-1; N2, nitrogen addition of 60 kg·hm-2·a-1. W0, no water addition; W1, 20% water addition; W2, 40% water addition.

图4 黑沙蒿种子功能性状相关性分析图。A, 常温下萌发性状与其余性状相关性热图。B, 低温下萌发性状与其余性状相关性热图。0.5 ab, 种子0.5 h吸水率; 1 ab, 种子1 h吸水率; 2 ab, 种子2 h吸水率; C, 种子碳含量; C:N, 种子碳氮含量比值; Cur, 种子曲率; FC, 种子形状系数; GE, 种子萌发势; GP, 种子萌发率; L, 种子长度; m, 种子质量; N, 种子氮含量; PA, 种子投影面积; S, 种子萌发速率; SA, 种子表面积; V, 种子体积; VI, 活力指数; W, 种子宽度; W:L, 种子长宽比。

Fig. 4 Correlation analyses among seed functional traits of Artemisia ordosica. A, Correlation map of germination traits at normal temperature with other traits. B, Correlation map of germination traits at low temperature with other traits. 0.5 ab, 0.5-hour water absorption of seeds; 1 ab, 1-hour water absorption of seeds; 2 ab, 2-hour water absorption; C, seed carbon content; C:N, ratio of seed carbon and nitrogen content; Cur, seed curvature; FC, seed form coefficient; GE, seed germination energy at normal temperature; GP, seed germination percentage at low temperature; L, seed length; m, seed mass; N, seed nitrogen content; PA, seed project area; S, seed germination speed at low temperature; SA, seed surface area; V, seed volume; VI, seed vigor index at normal temperature; W, seed width; W:L, ratio of seed width to length.

表3 黑沙蒿种子性状主成分(PC)分析结果

Table 3 Results of principal component (PC) analysis of seed traits of Artemisia ordosica

性状 Trait	对PC1的贡献度 Contribution to PC1 (%)	对PC2的贡献度 Contribution to PC2 (%)	对PC3的贡献度 Contribution to PC3 (%)	PC1的坐标值 Coordinate values in PC1	PC2的坐标值 Coordinate values in PC2	PC2的坐标值 Coordinate values in PC2
GP	19.62	0.31	0.61	0.87	0.08	0.08
S	12.82	0.98	29.46	0.71	-0.15	0.56
VI	12.51	5.80	20.92	0.70	-0.36	0.47
L	9.66	22.25	0.89	0.61	0.71	-0.10
N	6.74	13.22	0.08	0.51	-0.55	-0.03
m	0.16	28.07	1.08	0.08	0.80	0.11
Cur	0.63	23.22	1.82	-0.16	-0.73	-0.14
1 ab	3.85	5.34	21.23	-0.39	0.35	0.48
FC	16.97	0.03	6.90	-0.81	0.02	0.27
2 ab	17.05	0.78	17.02	-0.81	-0.13	0.43
方差占比 Variance percent (%)				38.86	22.73	10.77
累计方差占比 Cumulative variance percent (%)				38.86	61.59	72.36

图5 氮、水添加对黑沙蒿种子性状及萌发性状影响的主成分(PC)分析图。A, 黑沙蒿种子性状在两个主成分上的载荷。B, 黑沙蒿种子性状对氮添加的响应。C, 黑沙蒿种子性状对增水的响应。1 ab, 种子1 h吸水率; 2 ab, 种子2 h吸水率; Cur, 种子曲率; FC, 种子形状系数; GP, 常温下种子萌发率; L, 种子长度; m, 种子质量; N, 种子氮含量; S, 常温下种子萌发速率; VI, 常温下活力指数。N0, 不添加氮; N1, 氮添加20 kg·hm-2·a-1; N2, 氮添加60 kg·hm-2·a-1。W0, 不增水; W1, 增水20%; W2, 增水40%。

Fig. 5 Principal component (PC) analysis of the effects of nitrogen and water addition on seed traits and germination traits of Artemisia ordosica. A, Seed traits load of A. ordosica. B, Response of seed traits of A. ordosica to nitrogen increase. C, Response of seed traits of A. ordosica to water increase. 1 ab, 1-hour water absorption of seeds; 2 ab, 2-hour water absorption of seeds; Cur, seed curvature; GP, seed germination percentage at normal temperature; L, seed length; m, seed mass; N, seed nitrogen content; S, seed germination speed at normal temperature; VI, seed vigor index at normal temperature. N0, no nitrogen addition; N1, nitrogen addition of 20 kg·hm-2·a-1; N2, nitrogen addition of 60 kg·hm-2·a-1. W0, no water addition; W1, 20% water addition; W2, 40% water addition.

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氮水添加对黑沙蒿种子功能性状的影响

Effects of nitrogen and water addition on seed functional traits of Artemisia ordosica

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