不同放牧强度下内蒙古温带典型草原优势种植物防御策略

doi:10.17521/cjpe.2019.0329

植物生态学报 ›› 2020, Vol. 44 ›› Issue (6): 642-653.DOI: 10.17521/cjpe.2019.0329

不同放牧强度下内蒙古温带典型草原优势种植物防御策略

李颖¹, 龚吉蕊¹^,^*(), 刘敏¹, 侯向阳², 丁勇², 杨波¹, 张子荷¹, 王彪¹, 朱趁趁¹

¹北京师范大学地表过程与资源生态国家重点实验室, 中药资源保护与利用北京市重点实验室, 北京师范大学地理科学学部自然资源学院, 北京 100875
²中国农业科学院草地研究所, 呼和浩特 010021

收稿日期:2019-11-30 接受日期:2020-03-25 出版日期:2020-06-20 发布日期:2020-04-03
通讯作者: * 龚吉蕊: ORCID:0000-0003-2012-2606, jrgong@bnu.edu.cn
基金资助:
国家自然科学基金(41571048);国家重点研发计划(2016YFC0500502);国家重点基础研究发展计划(2014CB138803)

Defense strategies of dominant plants under different grazing intensity in the typical temperate steppe of Nei Mongol, China

LI Ying¹, GONG Ji-Rui¹^,^*(), LIU Min¹, HOU Xiang-Yang², DING Yong², YANG Bo¹, ZHANG Zi-He¹, WANG Biao¹, ZHU Chen-Chen¹

¹Laboratory of Earth Surface Processes and Resource Ecology, Beijing Key Laboratory of Traditional Chinese Medicine Protection and Utilization, School of Natural Resources, Faculty of Geographical Science, Beijing Normal University, Beijing 100875, China
²Grassland Research Institute of Chinese Academy of Agricultural Science, Hohhot 010021, China

Received:2019-11-30 Accepted:2020-03-25 Online:2020-06-20 Published:2020-04-03
Contact: GONG Ji-Rui: ORCID:0000-0003-2012-2606, jrgong@bnu.edu.cn
Supported by:
National Natural Science Foundation of China(41571048);National Key R&D Program of China(2016YFC0500502);State Key Basic R&D Plan of China(2014CB138803)

摘要/Abstract

摘要：

放牧是草原的主要利用方式, 但对牧草造成了一定的生物胁迫。面对生存压力, 牧草会通过调节初级与次级代谢过程启动防御机制。该研究以内蒙古温带典型草原优势种植物为研究对象, 通过测定其在5个放牧水平下不同营养器官中的次级代谢产物及木质素等含量, 探讨大针茅(Stipa grandis)和羊草(Leymus chinensis)的各营养器官在防御机制中的角色及其碳氮权衡策略。结果表明: 面对放牧胁迫, 大针茅和羊草会产生大量的单宁、总黄酮、酚类以及生物碱等次级代谢产物, 并以叶片为主要的合成及储存器官。中度放牧使羊草的化学防御机制得到较充分的诱导及表达。但二者地上部分木质素含量并无显著增加, 因此, 二者在避牧性机制上更倾向于化学防御而非机械防御。由于羊草有更高的氮利用效率, 这使得羊草可以通过碳氮两种代谢途径进行防御, 但大针茅在生长初期并不能将氮高效地分配到化学防御中。大针茅和羊草在生长初期面对轻度放牧胁迫, 较多的资源仍然用于初级代谢, 增加了植物耐牧性。因此, 轻度放牧有利于提高牧草的碳氮资源利用效率、提高生态系统生产力及稳定性。

关键词: 次级代谢, 防御机制, 碳氮权衡, 优势种, 放牧强度, 大针茅, 羊草

Abstract:

Aims Grazing is one of the main grassland using modes, which has caused certain biotic stress on pasture. In order to survive, pasture initiates defense mechanisms by regulating the primary and secondary metabolic processes. This article aims to 1) figure out the distribution of defensive substances in different organs produced by the dominant plants Stipa grandis and Leymus chinensis; 2) explore the similarities and differences of the defense mechanisms and carbon-nitrogen trade-off strategies in S. grandis and L. chinensis.
Methods We conducted a different grazing intensity experiment in the typical temperate steppe of Nei Mongol. The lignin and secondary metabolites, such as tannins, flavonoids, phenols and alkaloids, as well as the ratio of carbon and nitrogen in different organs of the dominant plants S. grandis and L. chinensis were investigated.
Important findings Our results showed that S. grandis and L. chinensis produced a large number of secondary metabolites such as tannins, flavonoids, phenols and alkaloids in the face of grazing stress. The leaves were the main synthesis and storage organs. In addition, the chemical defense mechanism of L. chinensis was well induced and expressed under moderate grazing. There was no significant increase in the lignin content in the aboveground parts of S. grandis and L. chinensis. Therefore, both of them were more inclined to chemical defense than mechanical defense in the mechanism of avoiding grazing. Because of the higher nitrogen use efficiency of L. chinensis, it could be protected by both carbon and nitrogen metabolic pathways. However, Stipa grandis did not efficiently distribute nitrogen into chemical defense in the early stages of growth. More resources still used for primary metabolism in the early stage of S. grandis and L. chinensis growth, under light grazing stress, which increased their grazing resistance. Therefore, light grazing is conducive to improving the carbon and nitrogen use efficiency of pasture, and improving ecosystem productivity and stability.

Key words: secondary metabolism, defense mechanism, carbon-nitrogen trade-off, dominant species, grazing intensity, Stipa grandis, Leymus chinensis

李颖, 龚吉蕊, 刘敏, 侯向阳, 丁勇, 杨波, 张子荷, 王彪, 朱趁趁. 不同放牧强度下内蒙古温带典型草原优势种植物防御策略. 植物生态学报, 2020, 44(6): 642-653. DOI: 10.17521/cjpe.2019.0329

LI Ying, GONG Ji-Rui, LIU Min, HOU Xiang-Yang, DING Yong, YANG Bo, ZHANG Zi-He, WANG Biao, ZHU Chen-Chen. Defense strategies of dominant plants under different grazing intensity in the typical temperate steppe of Nei Mongol, China. Chinese Journal of Plant Ecology, 2020, 44(6): 642-653. DOI: 10.17521/cjpe.2019.0329

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2019.0329

https://www.plant-ecology.com/CN/Y2020/V44/I6/642

图/表 8

图1 2016年内蒙古温带典型草原研究区月平均气温和降水量示意图。

Fig. 1 Monthly precipitation and mean air temperature at the study sites in the typical temperate steppe of Nei Mongol in 2016.

图2 6月(A、B)及8月(C、D)不同放牧水平下内蒙古温带典型草原大针茅与羊草的木质素含量(平均值±标准误差)。 CK, 围封; LG, 轻度放牧; MG, 中度放牧; HG, 重度放牧; EHG, 极重度放牧。不同小写字母表示处理间差异显著(p < 0.05)。

Fig. 2 Lignin content of Stipa grandis and Leymus chinensis in the typical temperate steppe of Nei Mongol in June (A, B) and August (C, D) under different grazing levels (mean ± SE). CK, grazing exclusion; LG, light grazing; MG, moderate grazing; HG, heavy grazing; EHG, extremely heavy grazing. Different lowercase letters indicate significant differences among treatments (p < 0.05).

图3 6月(A、B)及8月(C、D)不同放牧水平下内蒙古温带典型草原大针茅与羊草的单宁含量(平均值±标准误差)。 CK, 围封; LG, 轻度放牧; MG, 中度放牧; HG, 重度放牧; EHG, 极重度放牧。不同小写字母表示处理间差异显著(p < 0.05)。

Fig. 3 Tannins content of Stipa grandis and Leymus chinensis in the typical temperate steppe of Nei Mongol in June (A, B) and August (C, D) under different grazing levels (mean ± SE). CK, grazing exclusion; LG, light grazing; MG, moderate grazing; HG, heavy grazing; EHG, extremely heavy grazing. Different lowercase letters indicate significant differences among treatments (p < 0.05).

图4 6月(A、B)及8月(C、D)不同放牧水平下内蒙古温带典型草原大针茅与羊草的总黄酮含量(平均值±标准误差)。 CK, 围封; LG, 轻度放牧; MG, 中度放牧; HG, 重度放牧; EHG, 极重度放牧。不同小写字母表示处理间差异显著(p < 0.05)。

Fig. 4 Total flavonoids contents of Stipa grandis and Leymus chinensis in the typical temperate steppe of Nei Mongol in June (A, B) and August (C, D) under different grazing levels (mean ± SE). CK, grazing exclusion; LG, light grazing; MG, moderate grazing; HG, heavy grazing; EHG, extremely heavy grazing. Different lowercase letters indicate significant differences among treatments (p < 0.05).

图5 6月(A、B)及8月(C、D)不同放牧水平下内蒙古温带典型草原大针茅与羊草的总酚含量(平均值±标准误差)。 CK, 围封; LG, 轻度放牧; MG, 中度放牧; HG, 重度放牧; EHG, 极重度放牧。不同小写字母表示处理间差异显著(p < 0.05)。

Fig. 5 Total phenols contents of Stipa grandis and Leymus chinensis in the typical temperate steppe of Nei Mongol in June (A, B) and August (C, D) under different grazing levels (mean ± SE). CK, grazing exclusion; LG, light grazing; MG, moderate grazing; HG, heavy grazing; EHG, extremely heavy grazing. Different lowercase letters indicate significant differences among treatments (p < 0.05).

图6 6月及8月不同放牧水平下内蒙古温带典型草原大针茅与羊草叶片中的生物碱含量(平均值±标准误差)。 CK, 围封; LG, 轻度放牧; MG, 中度放牧; HG, 重度放牧; EHG, 极重度放牧。不同小写字母表示处理间差异显著(p < 0.05)。

Fig. 6 Alkaloid contents of Stipa grandis and Leymus chinensis in the typical temperate steppe of Nei Mongol in June and August under different grazing levels (mean ± SE). CK, grazing exclusion; LG, light grazing; MG, moderate grazing; HG, heavy grazing; EHG, extremely heavy grazing. Different lowercase letters indicate significant differences among treatments (p < 0.05).

图7 6月(A、B)及8月(C、D)不同放牧水平下内蒙古温带典型草原大针茅与羊草叶片碳氮比(平均值±标准误差)。 CK, 围封; LG, 轻度放牧; MG, 中度放牧; HG, 重度放牧; EHG, 极重度放牧。不同小写字母表示处理间差异显著(p < 0.05)。

Fig. 7 The C:N in leafs of Stipa grandis and Leymus chinensis in the typical temperate steppe of Nei Mongol in June (A, B) and August (C, D) under different grazing levels (mean ± SE). CK, grazing exclusion; LG, light grazing; MG, moderate grazing; HG, heavy grazing; EHG, extremely heavy grazing. Different lowercase letters indicate significant differences among treatments (p < 0.05).

表1 6月及8月内蒙古温带典型草原大针茅与羊草叶片中防御代谢产物与C:N的相关性分析

Table 1 Correlation analysis of defense metabolites and C:N in leaves of Stipa grandis and Leymus chinensis in the typical temperate steppe of Nei Mongol in June and August

		大针茅 S. grandis						羊草 L. chinensis
		单宁 Tannins	总黄酮 Flavonoids	总酚 Phenols	木质素 Lignin	总生物碱 Alkaloid	碳氮比 C:N	单宁 Tannins	总黄酮 Flavonoids	总酚 Phenols	木质素 Lignin	总生物碱 Alkaloid	碳氮比 C:N
大针茅 S. grandis	单宁 Tannins		0.800	0.900^*	-0.400	0.051	-0.300		0.400	0.400	-0.800	0.600	0.000
	总黄酮 Flavonoids			0.900^*	0.100	0.154	0.200			1.000^**	-0.200	-0.400	0.800
	总酚 Phenols				-0.300	0.051	-0.100				-0.200	-0.400	0.800
	木质素 Lignin					-0.154	0.500					-0.800	0.400
	总生物碱 Alkaloid						0.718						-0.800
	碳氮比 C:N
羊草 L. chinensis	单宁 Tannins		0.900^*	0.300	-0.200	0.600	0.500		0.000	0.600	-1.000^**	0.400	-0.600
	总黄酮 Flavonoids			0.600	-0.100	0.700	0.700			0.800	0.000	-0.800	-0.800
	总酚 Phenols				0.300	0.500	0.900^*				-0.600	-0.400	-1.000^**
	木质素 Lignin					0.600	-0.100					-0.400	0.600
	总生物碱 Alkaloid						0.300						0.400
	碳氮比 C:N

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不同放牧强度下内蒙古温带典型草原优势种植物防御策略

Defense strategies of dominant plants under different grazing intensity in the typical temperate steppe of Nei Mongol, China

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