Chin J Plan Ecolo ›› 2017, Vol. 41 ›› Issue (12): 1228-1238.doi: 10.17521/cjpe.2017.0115

• Research Articles • Previous Articles     Next Articles

Changes in leaf nitrogen and phosphorus stoichiometry of woody plants along an altitudinal gradient in Changbai Mountain, China

YANG Lei, SUN Han, FAN Yan-Wen, HAN Wei, ZENG Ling-Bing, LIU Chao, WANG Xiang-Ping*()   

  1. Key Laboratory for Forest Resources & Ecosystem Processes of Beijing, College of Forestry, Beijing Forestry University, Beijing 100083, China.
  • Online:2018-02-23 Published:2017-12-10
  • Contact: WANG Xiang-Ping E-mail:wangxiangping@bjfu.edu.cn

Abstract:

Aims Leaf is the organ of plant photosynthesis, and it is important to understand the drivers for the variations of leaf nitrogen (N) and phosphorus (P) stoichiometry along geographical and climatic gradients. Here we aimed to explore: 1) the changes in leaf nitrogen (N) and phosphorus (P) stoichiometry of woody plants along an altitudinal gradient in Changbai Mountain, and 2) the relative contribution of climate, plant characteristics, and phylogeny to the changes in leaf N, P concentration and N:P.

Methods We measured leaf N and P concentration, and N:P of 48 woody species in 14 plots along an altitudinal gradient in Changbai Mountain. General linear models (GLMs) and variation partitioning analyses were used to explain leaf N and P stoichiometry with parameters related to plant functional groups, climate and phylogenetic groups.

Important findings Leaf N concentration and N:P decreased with altitude, and were positively correlated with mean annual temperature (MAT) but negatively correlated with mean annual precipitation (MAP). Leaf P concentration had no significant relationship with altitude. Leaf N, P concentration and N:P were significantly different among different plant functional groups. Leaf N concentration of shrubs was higher than that of trees, but leaf P concentration and N:P did not differ significantly between shrubs and trees. Leaf N, P concentration and N:P of broad-leaved plants were significantly higher than those of coniferous plants. Similarly, leaf N, P concentration and N:P of deciduous plants were significantly higher than those of evergreen plants. The effects of climate on leaf N, P concentration and N:P were significant but very weak, with MAT and MAP together explaining 1.50%-2.98% of the variations. Phylogenetic groups explained 30.36%-54.38% of the variations, suggesting the critical effect of phylogeny. Our study also showed that the synergistic effect of climate and species composition (phylogeny) on leaf N and P stoichiometry was neglectable, at least along the altitude gradient of Changbai Mountain. More comparative studies between altitudinal and latitudinal gradients are needed to better understand the drivers of the changes in leaf stoichiometry along an environmental gradient.

Key words: leaf stoichiometry, altitude gradient, climate, plant functional groups, phylogeny, woody plants

Table 1

Statistics of leaf nitrogen (N), phosphorus (P) concentrations and N:P of the examined plant species in Changbai Mountain"

叶性状
Leaf trait
功能群
Functional group
样本量
n
平均值
Mean
最大值
Max
最小值
Min
标准偏差
SD
标准误差
SE
变异系数
CV
氮含量
N concentration (g·kg-1)
全部样品 All samples 431 21.46 37.90 5.10 6.75 0.33 0.31
生活型
Life form
乔木 Tree 240 20.74b 37.90 5.10 6.80 0.44 0.33
灌木 Shrub 191 22.37a 37.40 6.80 6.59 0.48 0.29
叶型
Leaf shape
针叶 Coniferous 89 15.50b 36.80 8.50 4.91 0.52 0.32
阔叶 Broad-leaved 342 23.02a 37.90 5.10 6.29 0.34 0.27
叶片习性
Leaf habit
常绿 Evergreen 87 14.49b 35.00 8.44 4.42 0.47 0.31
落叶 Deciduous 344 23.23a 37.90 5.10 6.06 0.33 0.26
磷含量
P concentration (g·kg-1)
全部样品 All samples 431 2.23 6.75 0.66 0.87 0.04 0.39
生活型
Life form
乔木 Tree 240 2.22 6.75 0.66 0.93 0.06 0.42
灌木 Shrub 191 2.25 5.29 0.77 0.80 0.06 0.35
叶型
Leaf shape
针叶 Coniferous 89 1.91b 4.13 0.66 0.77 0.08 0.40
阔叶 Broad-leaved 342 2.32a 6.75 0.77 0.88 0.05 0.38
叶片习性
Leaf habit
常绿 Evergreen 87 1.74b 3.30 0.66 0.66 0.07 0.38
落叶 Deciduous 344 2.36a 6.75 0.99 0.87 0.05 0.37
N:P 全部样品 All samples 431 10.81 33.70 2.47 4.77 0.23 0.44
生活型
Life form
乔木 Tree 240 10.60 33.70 2.47 4.89 0.32 0.46
灌木 Shrub 191 11.07 26.34 3.12 4.62 0.33 0.42
叶型
Leaf shape
针叶 Coniferous 89 9.35b 25.92 3.90 4.68 0.50 0.50
阔叶 Broad-leaved 342 11.19a 33.70 2.47 4.72 0.26 0.42
叶片习性
Leaf habit
常绿 Evergreen 87 9.75b 25.92 3.28 4.93 0.53 0.51
落叶 Deciduous 344 11.08a 33.70 2.47 4.70 0.25 0.42

Fig. 1

Leaf nitrogen (N) (A), phosphorus (P) (B) concentrations and N:P (C) in relation to altitude."

Fig. 2

Leaf nitrogen (N), phosphorus (P) concentrations and N:P in relation to mean annual temperature and mean annual precipitation."

Table 2

Contributions of each factor to the variations of leaf nitrogen (N), phosphorus (P) concentrations and N:P"

叶性状
Leaf traits
年平均气温
Mean annual
temperature
年降水量
Mean annual
precipitation
生活型
Life form
叶型
Leaf shape
叶片习性
Leaf habit
90 Mya谱系组
90 Mya division
60 Mya谱系组
60 Mya division
物种
Species
氮含量
N concentration
R2 0.012 0.012 0.016 0.229 0.324 0.415 0.459 0.536
p 0.023* 0.025* 0.008** 0.000*** 0.000*** 0.000*** 0.000*** 0.000***
磷含量
P concentration
R2 0.003 0.003 0.002 0.049 0.115 0.216 0.306 0.391
p 0.233 0.247 0.342 0.000*** 0.000*** 0.000*** 0.000*** 0.000***
N:P R2 0.016 0.016 0.003 0.028 0.018 0.152 0.215 0.303
p 0.008** 0.009** 0.257 0.000*** 0.005** 0.000*** 0.000*** 0.000***

Table 3

Summary of results of general linear model analysis for the effects of environmental factors, plant functional groups and phylogeny (90 and 60 Mya phylogenetic group, species) on leaf nitrogen (N), phosphorus (P) concentrations and N:P"

N P N:P
df %SS p df %SS p df %SS p
气候 Climate
年平均气温 Mean annual temperature 1 1.21 0.001** 1 0.33 0.146 1 1.65 0.002**
年降水量 Mean annual precipitation 1 0.61 0.022* 1 0.45 0.089 1 1.28 0.007**
植物功能型 Plant functional groups
生活型 Life form 1 1.54 0.000*** 1 0.22 0.237 1 0.27 0.214
叶型 Leaf shape 1 22.60 0.000*** 1 5.70 0.000*** 1 2.24 0.000***
叶片习性 Leaf habit 1 9.34 0.000*** 1 6.46 0.000*** 1 0.02 0.767
谱系 Phylogeny
90 Mya谱系组 90 Mya division 18 12.06 0.000*** 18 13.03 0.000*** 18 13.18 0.000***
60 Mya谱系组 60 Mya division 10 3.82 0.000*** 10 8.10 0.000*** 10 7.34 0.000***
物种 Species 17 5.01 0.000*** 17 6.33 0.002** 17 7.32 0.001**
残差 Residuals 380 43.81 380 59.36 380 66.72

Fig. 3

Variation partitioning analysis for leaf nitrogen (N), phosphorus (P) concentrations and N:P by three types of factors: Climate (mean annual temperature and mean annual precipitation), species (phylogenetic differences among species, He et al., 2010), and unexplained variations. A, leaf N concentration. B, leaf P concentration. C, N:P. a and b are the pure effects of climate and species, respectively; c is their synergistic effects. The significances of a and b were evaluated with F test. ***, p < 0.001; **, p < 0.01."

Appendix I

General information of the plots on Changbai Mountain"

样地 Plot 经度 Longitude (°E) 纬度 Latitude (°N) 海拔 Altitude (m) 林型 Forest type
CB01 128.11 42.39 530 白桦林 Betula platyphylla forest
CB02 128.08 42.41 650 阔叶红松林 Pinus koraiensis and broadleaf mixed forest
CB03 128.12 42.32 840 阔叶红松林 Pinus koraiensis and broadleaf mixed forest
CB04 128.17 42.23 950 白桦林 Betula platyphylla forest
CB05 128.17 42.23 970 阔叶红松林 Pinus koraiensis and broadleaf mixed forest
CB06 128.17 42.19 1 010 云冷杉林 Picea and Abies forest
CB07 128.11 42.12 1 270 阔叶红松林 Pinus koraiensis and broadleaf mixed forest
CB08 128.26 42.07 1 420 云冷杉林 Picea and Abies forest
CB09 128.09 42.10 1 420 长白落叶松林 Larix olgensis forest
CB10 128.24 42.08 1 440 长白落叶松林 Larix olgensis forest
CB11 128.08 42.09 1 530 长白落叶松林 Larix olgensis forest
CB12 128.07 42.07 1 660 长白落叶松林 Larix olgensis forest
CB13 128.07 42.06 1 885 岳桦林 Betula ermanii forest
CB14 128.07 42.06 1 940 岳桦林 Betula ermanii forest

Appendix II

Phylogenetic tree for 47 species in this study, based on the phylogenetic tree of Zanne et al. (2014). Showing the phylogenetic divisions at the 90 and 60 million years ago (Mya)"

Appendix III

Summary of general linear models for the effects of environmental variation (mean annual temperature and mean annual precipitation), plant functional groups and taxonomic variation (family, genus, and species) on individual leaf traits"

氮 Nitrogen 磷 Phosphorus N:P
df %SS p df %SS p df %SS p
气候 Climate
年平均气温 Mean annual temperature 1 1.21 0.001** 1 0.33 0.146 1 1.65 0.002**
年降水量 Mean annual precipitation 1 0.61 0.022* 1 0.45 0.089 1 1.28 0.007**
植物功能型 Plant functional groups
生活型 Life form 1 1.54 0.000*** 1 0.22 0.237 1 0.27 0.214
叶型 Leaf shape 1 22.60 0.000*** 1 5.70 0.000*** 1 2.24 0.000***
叶片习性 Leaf habit 1 9.34 0.000*** 1 6.46 0.000*** 1 0.02 0.767
系统发育关系 Taxonomy
科 Family 17 11.50 0.000*** 17 9.85 0.000*** 17 9.81 0.000***
属 Genus 10 3.11 0.003** 10 8.79 0.000*** 10 6.30 0.000***
种 Species 18 6.29 0.000*** 18 8.83 0.000*** 18 11.73 0.000***
残差 Residuals 380 43.81 380 59.36 380 66.72
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