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
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,*()
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:
YANG Wen-Gao1,ZI Hong-Biao1,CHEN Ke-Yu1,ADE Lu-Ji1,HU Lei1,WANG Xin1,WANG Gen-Xu2,WANG Chang-Ting1,*()
Received:
2018-12-31
Revised:
2019-04-02
Online:
2019-04-20
Published:
2019-08-29
Contact:
WANG Chang-Ting
Supported by:
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。
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
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).
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 ± 33.66bc | 3.85 ± 1.43ab | 0.84 ± 0.12a | 24.39 ± 4.24ab | 109.57 ± 39.04b | 4.58 ± 1.56bc |
毛白杨林 Populus tomentosa forest | 72.80 ± 14.33bc | 2.95 ± 0.52b | 0.75 ± 0.08a | 24.77 ± 2.83ab | 99.34 ± 25.78b | 3.99 ± 0.82c |
红桦林 Betula albosinensis forest | 133.81 ± 12.03a | 4.90 ± 0.77ab | 0.83 ± 0.02a | 27.53±2.20a | 161.04 ± 12.45a | 5.89 ± 0.82abc |
青扦林 Picea wilsonii forest | 99.11 ± 24.60ab | 3.49 ± 0.67ab | 0.85 ± 0.18a | 28.39 ± 4.14a | 118.21 ± 25.62ab | 4.23 ± 0.95bc |
山杨林 Populus davidiana forest | 59.21 ± 22.57c | 3.60 ± 0.94ab | 0.71 ± 0.16a | 16.6 ± 5.09c | 82.02 ± 18.60b | 5.29 ± 1.94abc |
圆柏林 Sabina chinensis forest | 88.97 ± 49.70bc | 5.86 ± 2.92a | 0.72 ± 0.81a | 15.70 ± 5.63c | 120.54 ± 59.85ab | 7.67 ± 3.53a |
云杉林 Picea asperata forest | 100.20 ± 37.05ab | 5.85 ± 2.80a | 0.87 ± 0.18a | 19.50 ± 7.88bc | 121.75 ± 44.38ab | 6.97 ± 3.06ab |
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 ± 33.66bc | 3.85 ± 1.43ab | 0.84 ± 0.12a | 24.39 ± 4.24ab | 109.57 ± 39.04b | 4.58 ± 1.56bc |
毛白杨林 Populus tomentosa forest | 72.80 ± 14.33bc | 2.95 ± 0.52b | 0.75 ± 0.08a | 24.77 ± 2.83ab | 99.34 ± 25.78b | 3.99 ± 0.82c |
红桦林 Betula albosinensis forest | 133.81 ± 12.03a | 4.90 ± 0.77ab | 0.83 ± 0.02a | 27.53±2.20a | 161.04 ± 12.45a | 5.89 ± 0.82abc |
青扦林 Picea wilsonii forest | 99.11 ± 24.60ab | 3.49 ± 0.67ab | 0.85 ± 0.18a | 28.39 ± 4.14a | 118.21 ± 25.62ab | 4.23 ± 0.95bc |
山杨林 Populus davidiana forest | 59.21 ± 22.57c | 3.60 ± 0.94ab | 0.71 ± 0.16a | 16.6 ± 5.09c | 82.02 ± 18.60b | 5.29 ± 1.94abc |
圆柏林 Sabina chinensis forest | 88.97 ± 49.70bc | 5.86 ± 2.92a | 0.72 ± 0.81a | 15.70 ± 5.63c | 120.54 ± 59.85ab | 7.67 ± 3.53a |
云杉林 Picea asperata forest | 100.20 ± 37.05ab | 5.85 ± 2.80a | 0.87 ± 0.18a | 19.50 ± 7.88bc | 121.75 ± 44.38ab | 6.97 ± 3.06ab |
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 |
器官 Organ | 项目 Item | 土壤 Soil | |||||
---|---|---|---|---|---|---|---|
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** |
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 | |||||
---|---|---|---|---|---|---|---|
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.
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 |
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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