青海森林生态系统中灌木层和土壤生态化学计量特征

doi:10.17521/cjpe.2018.0326

植物生态学报 ›› 2019, Vol. 43 ›› Issue (4): 352-364.DOI: 10.17521/cjpe.2018.0326

所属专题：生态化学计量

青海森林生态系统中灌木层和土壤生态化学计量特征

杨文高¹,字洪标¹,陈科宇¹,阿的鲁骥¹,胡雷¹,王鑫¹,王根绪²,王长庭^1,^*()

1 西南民族大学青藏高原研究院, 成都 610041
2 中国科学院成都山地灾害与环境研究所中国科学院山地表生过程与生态调控重点实验室, 成都 610041

收稿日期:2018-12-31 修回日期:2019-04-02 出版日期:2019-04-20 发布日期:2019-08-29
通讯作者: 王长庭
基金资助:
国家自然科学基金(31870407);国家自然科学基金(31370542);四川省重点研发项目(2018SZ0333);中央高校基本科研业务费专项资金(2018NZD13)

Ecological stoichiometric characteristics of shrubs and soils in different forest types in Qinghai, China

YANG Wen-Gao¹,ZI Hong-Biao¹,CHEN Ke-Yu¹,ADE Lu-Ji¹,HU Lei¹,WANG Xin¹,WANG Gen-Xu²,WANG Chang-Ting^1,^*()

1 Institute of Qinghai-Tibetan Plateau, Southwest University for Nationalities, Chengdu 610041, China
2 Key Laboratory of Mountain Surface Processes and Ecological Regulation of Chinese Academy of Sciences, Institute of Mountain Hazards and Environment, Chinese Academy of Sciences, Chengdu 610041, China

Received:2018-12-31 Revised:2019-04-02 Online:2019-04-20 Published:2019-08-29
Contact: WANG Chang-Ting
Supported by:
Supported by the Natural National Science Foundation of China(31870407);Supported by the Natural National Science Foundation of China(31370542);the Key R&D Projects in Sichuan Province(2018SZ0333);the Fundamental Research Funds for the Central Universities(2018NZD13)

摘要/Abstract

摘要：

灌木层作为森林生态系统的重要组成部分, 了解其生态化学计量特征将有助于揭示森林生态系统物质周转和养分循环等生态功能。该研究选取青海省7种主要优势林分——白桦(Betula platyphylla)林、毛白杨(Populus tomentosa)林、红桦(Betula albosinensis)林、青扦(Picea wilsonii)林、山杨(Populus davidiana)林、圆柏(Sabina chinensis)林、云杉(Picea asperata)林为研究对象, 采用野外取样和室内实验分析相结合的方法, 研究了不同林分林下灌木层不同器官(叶、枝干、根)及其表层(0-10 cm)土壤的碳(C)、氮(N)、磷(P)含量及其相关性。结果表明: 7种林分间灌木(叶、枝干、根) P含量、C:P均没有明显差异性; 山杨林、圆柏林、云杉林的林下灌木(叶、枝干、根) N含量、N:P高于白桦林、毛白杨林、红桦林和青扦林, C:N则相反。圆柏林的林下灌木生长受P限制, 其余6种林分的林下灌木生长受N限制。7种林分间土壤有机碳(SOC)和总氮(TN)含量呈现出明显差异性, 而总磷(TP)含量则差异不明显。相关性分析表明, 林下灌木(叶、枝干、根) N含量、C:N、N:P与土壤TN含量、C:N、N:P呈极显著相关性, 而P含量、C:P与土壤TP含量呈显著相关性。冗余分析表明, 林下灌木层植被C、N、P含量及生态化学计量特征受到土壤化学计量特征及各环境因子的共同影响, 其中土壤C:N、海拔、年平均气温、年降水量为主要影响因子。

关键词: 生态化学计量学, 森林生态系统, 林分类型, 灌木层, 环境因子, 青海

Abstract:

AimsAs a dominant understory layer, shrubs is important in the material turnover and nutrient circulation of forested ecosystems. It is essential to explore stoichiometric characteristics of carbon (C), nitrogen (N) and phosphorus (P) of the shrubs and their driving factors, including microenvironments and soil nutrients.
MethodsThe leaves, branches, roots of the shrubs and the soils they rooted were sampled from seven dominant forest types of Qinghai, China, and the tissue contents of C, N and P were examined. One-way ANOVA was used to explored the difference of the shrubs and the soils among the forest types using, respectively. Redundancy analysis (RDA) was used to analyze the effects of soils and environmental factors on the stoichiometric characteristics of C, N and P of shrubs.
Important findings Our results showed that there were no significant differences in the P content and C:P of the leaves, branches and roots among all the seven dominant forest types, while the N content and N:P of shrubs in the Populus davidiana, Sabina chinensis and Picea asperata forests were significantly higher than those in Betula platyphylla, Populus tomentosa, Betula albosinensis and Picea wilsonii forests, while the C:N ratios were the other way around. The shrubs in Sabina chinensis forest were limited by the soil P content, but that in the other six forest types was limited by the soil N content. The contents of soil organic C (SOC) and soil total N (TN) were significantly different among the seven forest types, while the soil total P (TP) was not. Correlation analysis showed that the N content, the C:N and N:P of understory shrub tissues (leaves, branches and roots) were significantly correlated with soil TN content, soil C:N and N:P, while tissue P contents and the C:P ratios were correlated with soil TP contents. Redundancy analysis (RDA) showed that the stoichiometric characteristics of C, N and P the understory shrub layer were synthetically affected by soils and environmental factors, of which the soil C:N, altitude, mean annual temperature and mean annual precipitation were the main influence factors.

Key words: ecological stoichiometry, forest ecosystem, forest type, shrub layer, environmental factor, Qinghai

杨文高, 字洪标, 陈科宇, 阿的鲁骥, 胡雷, 王鑫, 王根绪, 王长庭. 青海森林生态系统中灌木层和土壤生态化学计量特征. 植物生态学报, 2019, 43(4): 352-364. DOI: 10.17521/cjpe.2018.0326

YANG Wen-Gao, ZI Hong-Biao, CHEN Ke-Yu, ADE Lu-Ji, HU Lei, WANG Xin, WANG Gen-Xu, WANG Chang-Ting. Ecological stoichiometric characteristics of shrubs and soils in different forest types in Qinghai, China. Chinese Journal of Plant Ecology, 2019, 43(4): 352-364. DOI: 10.17521/cjpe.2018.0326

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2018.0326

https://www.plant-ecology.com/CN/Y2019/V43/I4/352

图/表 7

图1 青海森林样地点位图。

Fig. 1 Map of the sampling forests in Qinghai.

图2 青海不同林分林下灌木C、N、P含量特征(平均值±标准误差)。不同大写字母表示不同林分林下灌木相同器官C、N、P含量差异显著(p < 0.05), 不同小写字母表示相同林分林下灌木不同器官C、N、P含量差异显著(p < 0.05)。

Fig. 2 The C, N and P contents of shrubs of the different forest types in Qinghai (mean ± SE). Different uppercase letters indicate significant differences of the C, N and P contents in the same organs of shrubs under different forest types (p < 0.05), and different lowercase letters indicate significant differences of the C, N and P contents in different organs of shrubs under the same forest type (p < 0.05).

图3 青海不同林分林下灌木C:N、C:P、N:P特征(平均值±标准误差)。不同大写字母表示不同林分林下灌木相同器官C:N、C:P、N:P差异显著(p < 0.05), 不同小写字母表示相同林分林下灌木不同器官C:N、C:P、N:P差异显著(p < 0.05)。

Fig. 3 The C:N, C:P and N:P ratios in shrubs of different forest types in Qinghai (mean ± SE). Different uppercase letters indicate significant differences of the C:N、C:P、N:P in the same organs of shrubs under different forest types (p < 0.05), and different lowercase letters indicate significant differences of the C:N、C:P、N:P in different organs of shrubs under the same forest type (p < 0.05).

表1 青海不同林分土壤有机碳(SOC)、全氮(TN)、全磷(TP)含量及化学计量比特征(平均值±标准误差)

Table 1 Soil organic carbon (SOC), total nitrogen (TN), total phosphorus (TP) content and stoichiometry characteristics in different forest types in Qinghai (mean ± SE)

林分类型 Forest type	SOC含量 SOC content (mg·g^-1)	TN含量 TN content (mg·g^-1)	TP含量 TP content (mg·g^-1)	C:N	C:P	N:P
白桦林 Betula platyphylla forest	91.43 ± 33.66^bc	3.85 ± 1.43^ab	0.84 ± 0.12^a	24.39 ± 4.24^ab	109.57 ± 39.04^b	4.58 ± 1.56^bc
毛白杨林 Populus tomentosa forest	72.80 ± 14.33^bc	2.95 ± 0.52^b	0.75 ± 0.08^a	24.77 ± 2.83^ab	99.34 ± 25.78^b	3.99 ± 0.82^c
红桦林 Betula albosinensis forest	133.81 ± 12.03^a	4.90 ± 0.77^ab	0.83 ± 0.02^a	27.53±2.20^a	161.04 ± 12.45^a	5.89 ± 0.82^abc
青扦林 Picea wilsonii forest	99.11 ± 24.60^ab	3.49 ± 0.67^ab	0.85 ± 0.18^a	28.39 ± 4.14^a	118.21 ± 25.62^ab	4.23 ± 0.95^bc
山杨林 Populus davidiana forest	59.21 ± 22.57^c	3.60 ± 0.94^ab	0.71 ± 0.16^a	16.6 ± 5.09^c	82.02 ± 18.60^b	5.29 ± 1.94^abc
圆柏林 Sabina chinensis forest	88.97 ± 49.70^bc	5.86 ± 2.92^a	0.72 ± 0.81^a	15.70 ± 5.63^c	120.54 ± 59.85^ab	7.67 ± 3.53^a
云杉林 Picea asperata forest	100.20 ± 37.05^ab	5.85 ± 2.80^a	0.87 ± 0.18^a	19.50 ± 7.88^bc	121.75 ± 44.38^ab	6.97 ± 3.06^ab
平均值 Average	95.11 ± 37.63	5.17 ± 2.60	0.82 ± 0.16	20.55 ± 7.35	117.72 ± 44.32	6.35 ± 2.98

表2 青海不同林分灌木各器官与土壤C、N、P含量及化学计量比之间的相关性

Table 2 Correlation among the shrubs and soil C, N and P stoichiometric characteristics in different forest types in Qinghai

器官 Organ	项目 Item	土壤 Soil
器官 Organ	项目 Item	SOC	TN	TP	C:N	C:P	N:P
叶 Leaf	C	0.159^*	0.166^*	0.160^*	-0.078	0.100	0.106
	N	0.063	0.614^**	0.224^**	-0.759^**	-0.028	0.561^**
	P	0.005	0.214^**	0.427^**	-0.261^**	-0.185^*	0.022
	C:N	-0.054	-0.524^**	-0.134	0.662^**	0.001	-0.514^**
	C:P	-0.023	-0.116	-0.354^**	0.106	0.139	0.048
	N:P	0.067	0.496^**	-0.100	-0.617^**	0.122	0.596^**
枝干 Branch	C	0.048	0.012	0.032	-0.019	0.063	0.011
	N	-0.083	0.457^**	0.087	-0.742^**	-0.137	0.444^**
	P	-0.162^*	-0.005	0.230^**	-0.175^*	-0.277^**	-0.120
	C:N	0.026	-0.514^**	-0.068	0.757^**	0.072	-0.516^**
	C:P	0.122	0.012	-0.190^*	0.105	0.227^**	0.110
	N:P	0.018	0.328^**	-0.113	-0.453^**	0.063	0.397^**
根 Root	C	0.239^**	0.120	-0.084	0.142	0.301^**	0.175^*
	N	-0.084	0.340^**	-0.076	-0.575^**	-0.052	0.413^**
	P	-0.132	-0.024	0.188^*	-0.107	-0.209^**	-0.106
	C:N	0.058	-0.349^**	0.008	0.573^**	0.053	-0.396^**
	C:P	0.135	0.116	-0.176^*	-0.016	0.202^**	0.189^*
	N:P	-0.002	0.286^**	-0.207^**	-0.422^**	0.077	0.397^**

图4 青海不同林分灌木叶片(A)、枝干(B)和根(C) C、N、P含量及化学计量比与环境因子之间的冗余分析二维排序图。Altitude, 海拔; Canopy, 郁闭度; Slope, 坡度; MAP, 年降水量; MAT, 年平均气温; SC, 土壤碳含量; SN, 土壤氮含量; SP, 土壤磷含量; SCN, 土壤C:N; SCP, 土壤C:P; SNP, 土壤N:P; LC, 叶片碳含量; LN, 叶片氮含量; LP, 叶片磷含量; BC, 枝干碳含量; BN, 枝干氮含量; BP, 枝干磷含量; RC, 根碳含量; RN, 根氮含量; RP, 根磷含量。

Fig. 4 Two-dimensional sequence diagram of redundancy analysis between the C, N and P contents and their ratios in leaves (A), branches (B) and roots (C) of shrubs with environmental factors in different forest types in Qinghai. MAP, mean annual precipitation; MAT, mean annual air temperature; SC, soil carbon content; SN, soil nitrogen content; SP, soil phosphorus content; SCN, soil C:N; SCP, soil C:P; SNP, soil N:P; LC, leaf carbon content; LN, leaf nitrogen content; LP, leaf phosphorus content; BC, branch carbon content; BN, branch nitrogen content; BP, branch phosphorus content; RC, root carbon content; RN, root nitrogen content; RP, root phosphorus content.

表3 青海不同林分灌木环境因子解释量与显著性检验

Table 3 Explained variance of environmental factors and significant test of shrubs in different forest types in Qinghai

器官 Organ	环境因子 Environmental factors	环境因子解释量(%) Environmental factors explained variance	p	F
叶片 Leaf	SCN	44.2	0.002	132.5
	Altitude	8.0	0.002	32.68
	MAP	7.2	0.002	24.68
	MAT	2.5	0.002	10.94
	SNP	1.5	0.006	6.62
	SN	1.0	0.016	4.45
	SC	0.9	0.014	3.96
	SCP	0.7	0.052	3.22
	SP	0.5	0.128	2.29
	Slope	0.3	0.302	1.273
	Canopy	0.0	0.876	0.16
枝干 Branch	SCN	38.1	0.002	99.01
	Altitude	6.1	0.002	17.47
	MAP	3.9	0.002	12.09
	MAT	2.1	0.002	6.68
	SC	1.5	0.006	4.85
	SNP	1.2	0.018	4.03
	Canopy	0.7	0.104	2.25
	Slope	0.6	0.124	2.04
	SCP	0.4	0.288	1.19
	SP	0.3	0.434	0.84
	SN	0.1	0.638	0.37
根 Root	SCN	19.2	0.002	39.26
	MAT	5.2	0.002	12.31
	Altitude	4.2	0.002	8.98
	MAP	3.2	0.002	7.21
	SCP	3.1	0.002	7.16
	Canopy	1.3	0.046	3.14
	Slope	1.2	0.034	3.14
	SP	0.6	0.252	1.41
	SN	0.3	0.446	0.88
	SC	0.3	0.400	0.85
	SNP	0.2	0.538	0.63

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青海森林生态系统中灌木层和土壤生态化学计量特征

Ecological stoichiometric characteristics of shrubs and soils in different forest types in Qinghai, China

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