氮素富集对青藏高原东缘窄叶鲜卑花灌丛根系分泌物碳输入的影响

doi:10.17521/cjpe.2016.0329

植物生态学报 ›› 2017, Vol. 41 ›› Issue (6): 610-621.DOI: 10.17521/cjpe.2016.0329

所属专题：青藏高原植物生态学：植物-土壤-微生物

氮素富集对青藏高原东缘窄叶鲜卑花灌丛根系分泌物碳输入的影响

何为^1,², 杨雪英³, 肖娟⁴, 张子良^1,², 蒋铮^1,², 袁远爽^1,², 王东^1,², 刘庆¹, 尹华军^1,^4,^*()

¹中国科学院成都生物研究所, 中国科学院山地生态恢复与生物资源利用重点实验室, 生态恢复与生物多样性保育四川省重点实验室, 成都 610041
²中国科学院大学, 北京 100049
³四川省理县林业局, 四川理县 623100
⁴西华师范大学环境科学与工程学院, 四川南充 637000

收稿日期:2017-04-05 接受日期:2016-10-21 出版日期:2017-06-10 发布日期:2017-07-19
通讯作者: 尹华军
作者简介:* 通信作者Author for correspondence (E-mail:sunzhiqiang1956@sina.com)
基金资助:
基金项目中国科学院前沿科学重点研究项目(QYZDB-SSW-SMC023)、中国科学院青年创新促进会人才项目(20132420)、国家自然科学基金(31670449)和四川省青年科技基金(2016JQ0037)

Effects of nitrogen enrichment on root exudative carbon inputs in Sibiraea angustata shrubbery at the eastern fringe of Qinghai-Xizang Plateau

Wei HE^1,², Xue-Ying YANG³, Juan XIAO⁴, Zi-Liang ZHANG^1,², Zheng JIANG^1,², Yuan-Shuang YUAN^1,², Dong WANG^1,², Qing LIU¹, Hua-Jun YIN^1,^4,^*()

¹Key Laboratory of Mountain Ecological Restoration and Bioresource Utilization & Ecological Restoration Biodiversity Conservation Key Laboratory of Sichuan Province, Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610041, China;

²University of Chinese Academy of Sciences, Beijing 100049, China

³Lixian Forestry Bureau, Lixian, Sichuan 623100, China
and
⁴College of Environmental Science and Engineering, China West Normal University, Nanchong, Sichuan 637000, China

Received:2017-04-05 Accepted:2016-10-21 Online:2017-06-10 Published:2017-07-19
Contact: Hua-Jun YIN
About author:KANG Jing-yao(1991-), E-mail: kangjingyao_nj@163.com。

摘要/Abstract

摘要：

为探究高寒灌丛生态系统根系分泌物碳(C)输入通量对大气氮(N)沉降的响应规律, 该文以青藏高原东缘窄叶鲜卑花(Sibiraea angustata)灌丛为研究对象, 采用根系分泌物野外原位收集法, 分析了不同施N水平(对照N₀= 0 g·m^-2·a^-1; 低N处理N₅= 5 g·m^-2·a^-1; 高N处理N₁₀= 10 g·m^-2·a^-1)对根系分泌物C输入速率与通量季节动态变化规律的影响。结果表明: (1)窄叶鲜卑花灌丛单位根生物量、单位根长、单位根表面积根系分泌物C输入速率均表现出明显的季节性动态变化, 具体表现为8月> 6月> 10月, 并呈现出与5 cm土壤温度相一致的变化趋势。(2)施N降低了窄叶鲜卑花灌丛单位根生物量、单位根长和单位根表面积根系分泌物C输入速率, 但仅N₁₀处理与对照(N₀处理)间存在显著差异(p < 0.05)。(3) N₅和N₁₀处理下, 窄叶鲜卑花灌丛细根生物量与N₀处理相比分别降低了23.36%和33.84%。(4)由于施N导致根系分泌物C输入速率与细根生物量二者均显著降低, 使得施N对窄叶鲜卑花灌丛根系分泌物C输入通量(g·m^-2·a^-1)有显著的抑制作用, 并随着施N浓度的增加抑制作用增大。推测其可能的原因是N素富集在一定程度上缓和了植物根系对养分的微生物驱动需求, 从而降低了植物根系分泌物C输入通量, 即N素富集条件下植物采取了低N收益-低C投入的生理策略。该研究结果对于进一步认知不同环境变化下高寒灌丛生态系统根系分泌物C输入及其介导的土壤生物C-养分循环过程具有重要的理论意义。

关键词: 窄叶鲜卑花灌丛, 氮添加, 根系分泌物, 细根生物量, 根系分泌物通量

Abstract:

Aims Understanding the responses of root exudative carbon (C) to increasing nitrogen deposition is important for predicting carbon cycling in terrestrial ecosystems. However, fewer studies have investigated the dynamics of root exudation in shrubbery ecosystems compared to forests and grassland ecosystems. This objective of this study was to determine the effects of nitrogen fertilization on the rate and C flux of root exudates.Methods Three levels of nitrogen addition treatments were applied to a Sibiraea angustata shrubbery ecosystem situated at the eastern fringe of Qinghai-Xizang Plateau, including N₀(without nitrogen application), N₅(nitrogen addition rate of 5 g·m^-2·a^-1), and N₁₀(nitrogen addition rate of 10 g·m^-2·a^-1), respectively, in 5 m ´ 5 m plots. Root exudates were collected in June, August and October of 2015, using a modified culture-based cuvette system. Root biomass in each plot was measured with root core method.Important findings The rates of root exudates on biomass, length, and surface area basis all displayed apparent seasonal variations during the experimental period, with the magnitude ranked in the order of: August > June > October, consistent with changes in soil temperature at 5 cm depth. With increases in the nitrogen addition rate, the rate of root exudates on biomass, length, and area basis all trended lower. Compared with the control (N₀), the N₅ and N₁₀ treatments significantly reduced fine root biomass in the Sibiraea angustata shrubbery, by 23.36% and 33.84%, respectively. The decreasing root exudation and fine root biomass in response to nitrogen addition significantly decreased C flux of root exudates. Our results provide additional evidences toward a robust theoretical foundation for better understanding soil C-nutrient cycling process mediated by root exudation inputs in Alpine shrubbery ecosystems under various environmental changes.

Key words: Sibiraea angustata shrubbery, nitrogen addition, root exudation, root biomass, root exudation flux

何为, 杨雪英, 肖娟, 张子良, 蒋铮, 袁远爽, 王东, 刘庆, 尹华军. 氮素富集对青藏高原东缘窄叶鲜卑花灌丛根系分泌物碳输入的影响. 植物生态学报, 2017, 41(6): 610-621. DOI: 10.17521/cjpe.2016.0329

Wei HE, Xue-Ying YANG, Juan XIAO, Zi-Liang ZHANG, Zheng JIANG, Yuan-Shuang YUAN, Dong WANG, Qing LIU, Hua-Jun YIN. Effects of nitrogen enrichment on root exudative carbon inputs in Sibiraea angustata shrubbery at the eastern fringe of Qinghai-Xizang Plateau. Chinese Journal of Plant Ecology, 2017, 41(6): 610-621. DOI: 10.17521/cjpe.2016.0329

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2016.0329

https://www.plant-ecology.com/CN/Y2017/V41/I6/610

图/表 9

图1 根系分泌物取样装置结构示意图。A, 进样单元。B, 收集单元。C, 抽滤单元。1, 抽滤瓶; 2, 硅胶管; 3, 干燥管; 4, 真空泵。

Fig. 1 Schematic drawing of root exudate collection device. A, Sample injection unit. B, Collection unit. C, Suction filter unit. 1, suction flask; 2, silicone tube; 3, drying tube; 4, vacuum pump.

表1 不同研究结果植物根系分泌碳(C)输入速率比较与分析

Table 1 Comparison of root exudative carbon (C) input rates among studies

植物类型 Plant type	文献来源 Reference source	植物种 Plant species	单位根生物量根系分泌物C输入速率 Root exudative C input rate per root biomass (μg·g^-1·h^-1)	单位根长根系分泌物C输入速率 Root exudative C input rate per root length (μg·cm^-1·h^-1)	单位根表面积根系分泌物C输入速率 Root exudative C input rate per root surface area (μg·cm^-2·h^-1)
草本 Herbs	Personeni et al., 2007	玉米 Zea mays	-	-	2.10
	Li & Chen, 2011	美人蕉 Canna indica	175.42	-	-
		风车草 Cyperus flabelliformis	220.00	-	-
		水鬼蕉 Hymenocallis littoralis	16.79	-	-
	Cheng at al., 2016	小麦 Triticum aestivum	4.07-1.19	0.056-0.012
		平均值 Mean	83.50	0.03	2.10
乔木 Trees	Phillips et al., 2009	火炬松 Pinus taeda	2.00-40.00	-	-
	Phillips et al., 2011	火炬松 Pinus taeda	8.00-17.00	0.10-6.00	-
	Meier et al., 2013	火炬松 Pinus taeda	32.00	-	-
	Yin et al., 2013a	云杉 Picea asperata	-	0.70	9.30
	Yin et al., 2013a	冷杉 Abies faxoniana	-	0.50	4.90
	Yin et al., 2013b	云杉 Picea asperata	663.91-414.94	0.67-0.39	7.99-5.53
	Yin et al., 2014	北美鹅掌楸 Liriodendron tulipifera 糖枫 Acer saccharum	9.58	-	-
	Yin et al., 2014	北美红栎 Quercus rubra 美洲水青冈木 Fagus grandifolia	17.50	-	-
	Zhang et al., 2016	云杉 Picea asperata	396.76	-	-
	Li at al., 2014a	9年生云杉人工林 9-year-old Picea asperata	441.86	3.20	2.75
		13年生云杉人工林 13-year-old Picea asperata	284.88	3.66	1.01
		31年生云杉人工林 31-year-old Picea asperata	315.89	3.56	1.52
	Xiao, 2013	云杉幼苗 Picea asperata seedlings	79.16	0.70	9.30
	Xiao, 2013	冷杉幼苗 Abies faxoniana seedlings	65.55	0.50	4.90
	Qiao, 2015	云杉 Picea asperata	570.93	0.50	4.44
	Qiao, 2015	冷杉 Abies faxoniana	579.71	0.51	5.52
	Xiong et al., 2015	杉木 Cunninghamia lanceolata	173.68-138.04	0.37-0.14	1.67-0.46
	Xiong et al., 2015	米槠 Castanopsis carlesi	92.72-56.16	0.34-0.06	1.27-0.24
		平均值 Mean	203.07	1.29	4.05
灌木 Shrubs	本研究 This study	窄叶鲜卑花 Sibiraea angustata	26.22	0.02	0.29

表1 不同研究结果植物根系分泌碳(C)输入速率比较与分析

Table 1 Comparison of root exudative carbon (C) input rates among studies

植物类型 Plant type	文献来源 Reference source	植物种 Plant species	单位根生物量根系分泌物C输入速率 Root exudative C input rate per root biomass (μg·g^-1·h^-1)	单位根长根系分泌物C输入速率 Root exudative C input rate per root length (μg·cm^-1·h^-1)	单位根表面积根系分泌物C输入速率 Root exudative C input rate per root surface area (μg·cm^-2·h^-1)
草本 Herbs	Personeni et al., 2007	玉米 Zea mays	-	-	2.10
	Li & Chen, 2011	美人蕉 Canna indica	175.42	-	-
		风车草 Cyperus flabelliformis	220.00	-	-
		水鬼蕉 Hymenocallis littoralis	16.79	-	-
	Cheng at al., 2016	小麦 Triticum aestivum	4.07-1.19	0.056-0.012
		平均值 Mean	83.50	0.03	2.10
乔木 Trees	Phillips et al., 2009	火炬松 Pinus taeda	2.00-40.00	-	-
	Phillips et al., 2011	火炬松 Pinus taeda	8.00-17.00	0.10-6.00	-
	Meier et al., 2013	火炬松 Pinus taeda	32.00	-	-
	Yin et al., 2013a	云杉 Picea asperata	-	0.70	9.30
	Yin et al., 2013a	冷杉 Abies faxoniana	-	0.50	4.90
	Yin et al., 2013b	云杉 Picea asperata	663.91-414.94	0.67-0.39	7.99-5.53
	Yin et al., 2014	北美鹅掌楸 Liriodendron tulipifera 糖枫 Acer saccharum	9.58	-	-
	Yin et al., 2014	北美红栎 Quercus rubra 美洲水青冈木 Fagus grandifolia	17.50	-	-
	Zhang et al., 2016	云杉 Picea asperata	396.76	-	-
	Li at al., 2014a	9年生云杉人工林 9-year-old Picea asperata	441.86	3.20	2.75
		13年生云杉人工林 13-year-old Picea asperata	284.88	3.66	1.01
		31年生云杉人工林 31-year-old Picea asperata	315.89	3.56	1.52
	Xiao, 2013	云杉幼苗 Picea asperata seedlings	79.16	0.70	9.30
	Xiao, 2013	冷杉幼苗 Abies faxoniana seedlings	65.55	0.50	4.90
	Qiao, 2015	云杉 Picea asperata	570.93	0.50	4.44
	Qiao, 2015	冷杉 Abies faxoniana	579.71	0.51	5.52
	Xiong et al., 2015	杉木 Cunninghamia lanceolata	173.68-138.04	0.37-0.14	1.67-0.46
	Xiong et al., 2015	米槠 Castanopsis carlesi	92.72-56.16	0.34-0.06	1.27-0.24
		平均值 Mean	203.07	1.29	4.05
灌木 Shrubs	本研究 This study	窄叶鲜卑花 Sibiraea angustata	26.22	0.02	0.29

图2 不同施氮(N)处理下窄叶鲜卑花灌丛根系分泌物碳(C)输入速率差异(平均值±标准偏差, n = 3)。A, 单位根生物量根系分泌物C输入速率。B, 单位根长根系分泌物C输入速率。C, 单位根表面积根系分泌物C输入速率。不同小写字母表示同一采样时间各处理间在p < 0.05水平上差异显著。N0、N5和N10为3种施氮梯度(N0: 0 g·m-2·a-1; N5: 5 g·m-2·a-1; N10: 10 g·m-2·a-1)。

Fig. 2 Differences in root exudative carbon (C) input rates in Sibiraea angustata shrubbery among different nitrogen fertilization treatments (mean ± SD, n = 3). A, Root exudative C input rate per root biomass. B, Root exudative C input rate per root length. C, Root exudative C input rate per root surface area. Different lowercase letters indicate significant differences (p < 0.05) among treatments on a given sampling date. Three nitrogen addition levels: N0 (0 g·m-2·a-1), N5 (5 g·m-2·a-1), N10 (10 g·m-2·a-1).

表2 窄叶鲜卑花灌丛根系分泌物碳(C)输入速率对施氮(N)和采样日期的重复测量方差分析p值

Table 2 Summary of repeated measures ANOVA showing the p values for responses of root exudative carbon (C) input rates of Sibiraea angustata shrubbery to nitrogen (N) fertilization and sampling date

	施N N fertilization	采样日期 Sampling date	采样日期×施N Sampling date × N fertilization
单位根生物量根系分泌物C输入速率 Root exudative C input rates per root biomass	<0.001	<0.001	0.255
单位根长根系分泌物C输入速率 Root exudative C input rates per root length	<0.001	0.019	0.844
单位根表面积根系分泌物C输入速率 Root exudative C input rates per root surface area	<0.001	<0.001	<0.001

图3 窄叶鲜卑花灌丛单位根生物量根系分泌物碳(C)输入速率与5 cm土壤温度的相关性分析。N0、N5和N10为3种施氮梯度(N0: 0 g·m-2·a-1; N5: 5 g·m-2·a-1; N10: 10 g·m-2·a-1)。

Fig. 3 Correlation analysis between root exudative carbon (C) input rate per root biomass and soil temperature at 5 cm depth. Three nitrogen addition levels: N0 (0 g·m-2·a-1), N5 (5 g·m-2·a-1), N10 (10 g·m-2·a-1).

图4 不同施N处理下窄叶鲜卑花灌丛细根生物量变化(平均值±标准偏差, n = 3)。不同小写字母表示同一采样时间各处理间在p < 0.05水平上差异显著。N0、N5和N10为3种施氮梯度(N0: 0 g·m-2·a-1; N5: 5 g·m-2·a-1; N10: 10 g·m-2·a-1)。

Fig. 4 Changes in fine root biomass in Sibiraea angustata shrubbery under different nitrogen fertilization treatments (mean ± SD, n = 3). Different lowercase letters indicate significant differences among treatments on a given sampling date (p < 0.05). Three nitrogen addition levels: N0 (0 g·m-2·a-1), N5 (5 g·m-2·a-1), N10 (10 g·m-2·a-1).

表3 不同施氮(N)处理下窄叶鲜卑花灌丛根系分泌物碳(C)通量(平均值±标准偏差, n = 3)

Table 3 Root exudative carbon (C) flux in Sibiraea angustata shrubbery under different nitrogen (N) fertilization treatments (mean ± SD, n = 3)

处理 Treatment	单位根生物量根系分泌物C输入速率 Root exudative C input rate per root biomass (mg·g^-1·d^-1)	细根生物量 Fine root biomass (g·m^-2)	根系分泌物C通量 C flux of root exudates (g·m^-2·a^-1)
N₀	0.68 ± 0.02^a	169.17 ± 47.83^a	16.89 ± 0.31^a
N₅	0.57 ± 0.04^b	129.65 ± 81.71^b	9.58 ± 0.24^b
N₁₀	0.36 ± 0.02^c	111.93 ± 58.49^c	5.81 ± 0.12^c

表4 不同施N水平下土壤NO3--N、NH4+-N含量(平均值±标准偏差, n =3)

Table 4 Changes in nitrate nitrogen and ammonium nitrogen under different nitrogen fertilization treatments (mean ± SD, n = 3)

处理 Treatment	2015-06		2015-08			2015-10
处理 Treatment	NO₃^--N (mg·kg^-1)	NH₄⁺-N (mg·kg^-1)	NO₃^--N (mg·kg^-1)	NH₄⁺-N (mg·kg^-1)		NO₃^--N (mg·kg^-1)	NH₄⁺-N (mg·kg^-1)
N₀	20.88 ± 0.91^a	7.44 ± 0.16^a	23.92 ± 0.23^a	8.66 ± 0.54^a	25.88 ± 0.83^a		7.63 ± 0.28^a
N₅	22.19 ± 0.71^a	12.57 ± 0.64^b	24.51 ± 0.31^a	13.21 ± 0.27^b	26.21 ± 0.24^a		12.42 ± 0.59^b
N₁₀	30.38 ± 0.71^b	14.29 ± 0.85^b	28.98 ± 0.65^b	16.18 ± 0.56^b	29.38 ± 0.17^b		13.94 ± 0.71^b

图5 不同施N处理下土壤微生物磷脂脂肪酸(PLFAs)总量变化(平均值±标准偏差, n = 3)。不同小写字母表示同一采样时间各处理间在p < 0.05水平上差异显著。N0、N5和N10为3种施氮梯度(N0: 0 g·m-2·a-1; N5: 5 g·m-2·a-1; N10: 10 g·m-2·a-1)。

Fig. 5 Changes in total soil microbial phospholipid fatty acid (PLFAs) under different nitrogen fertilization treatments (mean ± SD, n = 3). Different lowercase letters indicate significant differences (p < 0.05) among treatments on a given sampling date. Three nitrogen addition levels: N0 (0 g·m-2·a-1), N5 (5 g·m-2·a-1), N10 (10 g·m-2·a-1).

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氮素富集对青藏高原东缘窄叶鲜卑花灌丛根系分泌物碳输入的影响

Effects of nitrogen enrichment on root exudative carbon inputs in Sibiraea angustata shrubbery at the eastern fringe of Qinghai-Xizang Plateau

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