Stoichiometric characteristics of carbon and nitrogen in plants and their influencing factors in the lower reaches of the Heihe River, northwestern China
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摘要:目的 碳 、氮元素对植物的生长发育至关重要，尤其是在极端干旱的内陆河流域，不同功能群植物对土壤水分和养分需求不同，往往表现出不同的化学计量特征。探寻黑河下游不同功能群植物的化学计量特征及对地下水变化的养分反应，对进一步了解全球气候变化背景下荒漠生态系统具有重要意义。方法 在黑河下游，根据与河道的距离，共设置22个采样点，采用相关性分析、方差分解方法分别对不同功能群植物的碳氮化学计量特征及其与环境因子的关系进行研究。结果 研究结果表明：黑河下游地区植物叶片和细根的碳元素含量平均值分别为408.53和16.30 mg/g；氮元素含量平均值分别为500.34和11.81 mg/g，碳氮比平均值分别为30.74和49.48。与全球和区域尺度研究相比，黑河下游植物具有较高的细根碳含量、较低的氮含量和较高的碳氮比。在不同地下水位梯度上，木本植物的碳含量、氮含量与碳氮比均与地下水深度显著相关，但草本植物与地下水不相关。黑河下游植物碳氮化学计量特征与土壤理化属性紧密相关。地下水和土壤含水量、土壤总氮共同解释了木本植物化学计量总变异的53% ~ 75%；土壤pH值和土壤电导率共同解释了草本植物化学计量总变异的20%。结论 面对极端干旱盐碱环境，水分是影响木本植物碳氮化学计量特征变化的关键因子，草本植物碳氮化学计量特征主要受土壤pH值和土壤电导率影响。Abstract:Objective Carbon (C) and nitrogen (N) elements are crucial for plant growth and development, especially in extremely arid inland river basins. Plants in different functional groups would show varied stoichiometric characteristics due to the variations in requirements for soil water and nutrients. It is of great significance to explore stoichiometric characteristics among different plant functional groups in the lower reaches of the Heihe River of northwestern China, as well as their nutrient contents responsing to groundwater fluctuation, aiming to further understand desert ecosystems under the background of global climate change.Method In the lower reaches of the Heihe River, a total of 22 sampling sites were set up according to the vertical distance between vegetation and river. Correlation analysis and variation partition analysis (VPA) were applied to determine the relationship between plant stoichiometry and environmental factors among plant functional groups, respectively.Result The average C contents of leaves and fine roots of plants in the lower reaches of Heihe River were 408.53 and 16.30 mg/g, the average N contents were 500.34 and 11.81 mg/g, and the average C∶N ratios were 30.74 and 49.48, respectively. Compared with global and regional studies, it was found that plants in the lower reaches of the Heihe River had higher C content, lower N content and higher C∶N. Different from herbaceous plants, the C content, N content and C∶N of woody plants were significantly correlated with the changes of groundwater depth. We found that the stoichiometric characteristics of plant carbon and nitrogen in the lower reaches of the Heihe River were significantly correlated with soil properties. The groundwater and soil variables jointly explained 53%−75% of the variation in woody plant stoichiometry. Additionally, soil pH and soil electrical conductivity explained 20% of the variation in herbaceous plant stoichiometry.Conclusion Our study finds that groundwater is the key factor influencing the carbon and nitrogen stoichiometry of woody plants under extreme drought and saline-alkali environment, and the carbon and nitrogen stoichiometry of herbaceous plants is mainly influenced by soil pH and soil electrical conductivity.
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Keywords:
- groundwater /
- Heihe River /
- carbon and nitrogen /
- stoichiometric characteristics /
- functional group
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图 2 木本植物碳氮化学计量特征与地下水深度关系
Figure 2. Relationship between carbon and nitrogen stoichiometric characteristics of woody plants and underground water depth
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图 3 草本植物碳氮化学计量特征与地下水深度关系
Figure 3. Relationship between carbon and nitrogen stoichiometric characteristics of herbaceous plants and underground water depth
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图 4 环境因子对木本植物叶片和细根的碳、氮化学计量特征的解释
GW.地下水；SM1.土壤0 ~ 20 cm含水量；SM2.土壤20 ~ 40 cm含水量；SM4.土壤60 ~ 100 cm含水量；SEC.土壤电导率；STN.土壤全氮含量；NE.未解释率。NL.叶片氮含量；CL.叶片碳含量；CNL.叶片碳氮比；NR.细根氮含量；CR.细根碳含量；CNR.细根碳氮比。下同。GW, underground water; SM1, 0−20 cm soil water content; SM2, 20−40 cm soil water content; SM4, 60−100 cm soil water content; SEC, soil electrical conductivity; STN, soil total nitrogen content; NE, no-explanation rate. NL, leaf N concentration; CL, leaf C concentration; CNL, C∶N of leaf; NR, fine root N concentration; CR, fine root C concentration; CNR, C∶N of fine root. The same below.
Figure 4. Explanation of environmental factors on the carbon and nitrogen stoichiometric characteristics of leaves and fine roots of woody plants
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图 5 环境因子对草本植物叶片和细根的碳、氮化学计量特征的解释
pH.土壤pH；SEC.土壤电导率。pH, soil pH；SEC, soil electrical conductivity.
Figure 5. Explanation of environmental factors on the carbon and nitrogen stoichiometric characteristics of herbaceous leaves and fine roots
表 1 采样点基本信息
Table 1 Basic information of the sampling sites
样点
Sample site地下水深度
Underground water depth/m木本植物 Woody plant 草本植物 Herb 优势种
Dominant species重要值
Importance value优势种
Dominant species重要值
Importance valueS1 1.46 多枝柽柳 Tamarix ramosissima 0.53 S2 1.51 胡杨 Populus euphratica 0.58 苦豆子 Sophora alopecuroides 0.37 S3 1.77 胡杨 Populus euphratica 0.63 苦豆子 Sophora alopecuroides 0.21 S4 1.78 胡杨 Populus euphratica 0.48 胀果甘草 Glycyrrhiza inflata 0.14 S5 1.96 多枝柽柳 Tamarix ramosissima 0.68 骆驼蓬 Peganum harmala 0.32 S6 2.00 黑果枸杞 Lycium ruthenicum 0.23 苦豆子 Sophora alopecuroides 0.49 S7 2.08 胡杨 Populus euphratica 0.48 芨芨草 Achnatherum splendens 0.42 S8 2.26 胡杨 Populus euphratica 0.54 苦豆子 Sophora alopecuroides 0.44 S9 2.30 多枝柽柳 Tamarix ramosissima 0.35 花花柴 Karelinia caspia 0.39 S10 2.49 白刺 Nitraria tangutorum 0.33 沙蒿 Artemisia desertorum 0.38 S11 2.56 多枝柽柳 Tamarix ramosissima 0.24 苦豆子 Sophora alopecuroides 0.70 S12 2.73 红砂 Reaumuria soongorica 0.80 沙蒿 Artemisia desertorum 0.10 S13 2.79 膜果麻黄 Ephedra przewalskii 0.43 驼蹄瓣 Zygophyllum fabago 0.14 S14 2.85 骆驼刺 Alhagi sparsifolia 0.70 花花柴 Karelinia caspia 0.38 S15 2.89 红砂 Reaumuria soongorica 0.37 花花柴 Karelinia caspia 0.54 S16 3.00 多枝柽柳 Tamarix ramosissima 0.83 骆驼蓬 Peganum harmala 0.26 S17 3.15 膜果麻黄 Ephedra przewalskii 0.85 S18 3.30 红砂 Reaumuria soongorica 0.71 骆驼蓬 Peganum harmala 0.12 S19 3.31 膜果麻黄 Ephedra przewalskii 0.47 S20 3.44 膜果麻黄 Ephedra przewalskii 0.54 S21 3.55 膜果麻黄 Ephedra przewalskii 0.85 S22 3.66 膜果麻黄 Ephedra przewalskii 0.66 
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表 2 不同功能群植物的叶片和细根养分组成
Table 2 Nutrient composition of leaves and fine roots of different functional groups of plants
指标 Index 总体 Total 木本植物 Woody plant 草本植物 Herb 叶片氮含量 Leaf N concentration/(mg·g−1) 16.30 ± 0.59 15.06 ± 0.75b 18.66 ± 1.02a 叶片碳含量 Leaf C concentration/(mg·g−1) 408.54 ± 10.89 396.80 ± 13.63a 429.05 ± 17.41a 叶片碳氮比 Leaf C∶N 30.74 ± 1.44 31.69 ± 1.83a 29.30 ± 2.54a 细根氮含量 Fine root N concentration/(mg·g−1) 11.81 ± 0.44 12.61 ± 0.47a 10.42 ± 1.03b 细根碳含量 Fine root C concentration/(mg·g−1) 500.34 ± 15.10 514.52 ± 18.92a 484.33 ± 26.14a 细根碳氮比 Fine root C∶N 49.49 ± 2.69 45.35 ± 3.09b 54.27 ± 5.14a 注：同行不同字母表示差异显著（P < 0.05）。 Note: different letters in the same line indicate significant difference (P < 0.05).
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表 3 土壤因子对不同功能群植物化学计量的影响
Table 3 Effects of soil factors on plant stoichiometry of different functional groups
分类
Classification指标
Index电导率
Electrical conductivity土壤pH
Soil pH0 ~ 20 cm
含水量
0−20 cm
water content20 ~ 40 cm
含水量
20−40 cm
water content40 ~ 60 cm
含水量
40−60 cm
water content60 ~ 100 cm
含水量
60−100 cm
water content土壤总碳
Soil total carbon土壤全N
Soil total nitrogen土壤全P
Soil total phosphorus木本植物
Woody plant叶片氮含量
Leaf N concentration−0.10 0.14 0.50** 0.48** 0.37** 0.39** 0.27* 0.21** 0.13 叶片碳含量
Leaf C concentration0.03 0.04 −0.01 −0.05 −0.04 −0.24* −0.03 0.20 0.11 叶片碳氮比
Leaf C∶N0.30* −0.17 −0.28* −0.28* −0.25* −0.35** −0.15 −0.01 0.04 细根氮含量
Fine root N concentration0.25 −0.32 −0.41** −0.37** −0.34** −0.39** −0.37** −0.21** 0.19 细根碳含量
Fine root C concentration−0.07 −0.06 −0.34** −0.43** −0.45** −0.49** −0.27* −0.17 0.17 细根碳氮比
Fine root C∶N−0.27* 0.27* 0.21 0.11 0.09 0.03 0.18 0.11 −0.05 草本植物
Herb叶片氮含量
Leaf N concentration0.44* −0.40* −0.24 −0.30 −0.07 −0.18 −0.05 0.36 0.14 叶片碳含量
Leaf C concentration−0.17 0.26 0.46* 0.13 0.24 0.32 0.23 0.01 −0.16 叶片碳氮比
Leaf C∶N−0.38* 0.44* 0.32 0.18 0.04 0.20 0.02 −0.31 −0.26 细根氮含量
Fine root N concentration0.42* −0.45* −0.29 −0.25 −0.20 −0.29 −0.21 0.07 0.58 细根碳含量
Fine root C concentration−0.16 0.25 0.14 0.11 0.04 0.13 0.15 0.13 0.09 细根碳氮比
Fine root C∶N−0.38* 0.48** 0.21 −0.01 0.00 0.13 0.12 0.02 −0.10 注：* 表示相关性显著（P < 0.05）；**表示相关性极显著（P < 0.01）．Notes: * indicates significant correlation at P < 0.05 level; ** indicates extremely significant correlation at P < 0.01 level．
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