Leaf stoichiometry of two conifer species with different life forms in northern Daxing’an Mountains
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摘要:
目的 研究寒温带落叶树种和常绿树种针叶生态化学计量特征的差异及随龄级增加的变化规律，揭示不同生活型针叶树种生态化学计量特征与生长阶段之间的关系，为寒温带植物生长与养分供给的研究提供帮助。 方法 以大兴安岭北部兴安落叶松和樟子松为对象，研究5—9月针叶C、N、P含量及化学计量特征，利用内稳性指数和Pearson相关系数分析C、N、P之间及与环境因子的相关性。 结果 （1）不同生长阶段兴安落叶松和樟子松针叶N、P月平均含量表现为随龄级的增长而降低，C含量则是随龄级的增加而增加，且5—8月兴安落叶松针叶N、P含量高于樟子松，C含量则低于樟子松。（2）兴安落叶松针叶C∶N、C∶P和N∶P月平均值均高于樟子松，4个龄级兴安落叶松针叶N∶P均小于14，而樟子松在5—7月N∶P小于14，8—9月N∶P大于16。兴安落叶松针叶N、P以及N∶P稳定性大于樟子松。（3）土壤含水率与2树种针叶生态化学计量特征呈极显著相关，樟子松针叶N含量与C、N、P化学计量比呈显著相关，兴安落叶松针叶C含量与N∶P、土壤铵态氮和硝态氮显著相关。 结论 大兴安岭北部不同生活型的2种针叶树种，叶片生态化学计量对寒冷气候和冻土生境的适应策略不同，兴安落叶松是本区的顶级群落优势种，但其生长受到N的限制，而樟子松生长的限制因子因季节变化而不同。 Abstract:Objective Study on the difference in ecological stoichiometric characteristics and their changes with increasing age of conifers of deciduous and evergreen species in the cold temperate zone to reveal the relationship between ecological stoichiometric characteristics and growth stage of two different life forms of conifers, and to provide help for the study of plant growth and nutrient supply in cold-temperate of China. Method The contents and stoichiometric characteristics of C, N, P in Larix gmelinii and Pinus sylvestris var. mongolica from May to September were studied. The correlation between C, N, P and environmental factors was analyzed by homeostasis index and Pearson correlation coefficient. Result (1) During the different growth stage, monthly average N, P content of Larix gmelinii and Pinus sylvestris var. mongolica needles were both decreased with the increase of age class, while the C content increased with the increase of age class. The N and P contents of needles of Larix gmelinii from May to August were higher than those of Pinus sylvestris var. mongolica, while the C content was lower than those of Pinus sylvestris var. mongolica. (2) The monthly mean values of C∶N, C∶P and N∶P of needles of Larix gmelinii were higher than those of Pinus sylvestris var. mongolica. The N∶P of needles of Larix gmelinii of four age classs were all less than 14, while the N∶P of needles of Pinus sylvestris of four age classes from May to July was less than 14, and the N∶P of needles of Pinus sylvestris var. mongolica of four age classes was more than 16. The homeostasis of N, P and N∶P of Larix gmelinii needles was greater than that of Pinus sylvestris var. mongolica. (3) Soil moisture content was significantly correlated with the ecological stoichiometric characteristics of the needles of the two species. The N content of needles of Pinus sylvestris var. mongolica was significantly correlated with the stoichiometric ratio of C, N and P, and the C content of needles of Larix gmelinii was significantly correlated with N∶P and soil ammonium and nitrate nitrogen. Conclusion The two conifer species with different life forms in the northern Daxing’an Mountains have different adaptation strategies to cold climate and frozen soil.The Larix gmelinii is the dominant species in the top community, but its different growth stages were both limited by N. However, Pinus sylvestris var. mongolica the growth limiting factors of Pinus sylvestris var. mongolica are different due to seasonal changes. -
Key words:
- cold temperate zone /
- nutrient content /
- Stoichiometry /
- Conifer species /
- Age class
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图 1 兴安落叶松林和樟子松林树龄与胸径关系图
Figure 1. Relationship between tree Age and DBH of Larix gmelinii forest and Pinus sylvestris var. mongolica forest
图 2 不同龄级兴安落叶松和樟子松针叶氮含量变化特征
同一月份不同龄级间差异用大写字母表示A、B、C；同一龄级不同月份间差异用小写字母表示a、b、c。下同。Differences between different age groups in the same month are represented by capital letters A, B and C; Differences between the same age group and different months are represented by lowercase letters a, b and c. The same below.
Figure 2. Variation characteristics of nitrogen content in needles of Larix gmelinii and Pinus sylvestris var. mongolica at different age levels
图 3 不同龄级兴安落叶松和樟子松针叶磷含量变化特征
Figure 3. Phosphorus content changes in needles of Larix gmelinii and Pinus sylvestris var. mongolica at different age levels
图 4 不同龄级兴安落叶松和樟子松针叶碳含量变化特征
Figure 4. Variation characteristics of carbon content in needles of Larix gmelinii and Pinus sylvestris var. mongolica at different age levels
图 5 不同龄级兴安落叶松和落叶松针叶碳氮磷化学计量比变化特征
Figure 5. C, N and P stoichiometric ratios of Larix gmelinii and Pinus sylvestris var. mongolica needles at different age levels
表 1 样地基本情况
Table 1. Basic situation of the sample site
森林植被类型
Forest vegetation types海拔
Altitude/m坡度
Gradient/（°）平均胸径
DBH/cm平均树高
Hight of tree/m树种组成
Species composition密度/（株·hm−2）
Density/（tree·ha−1）郁闭度
Canopy density兴安落叶松林
Larix gmelinii forest305 5 13.67 11.44 8落1樟1白 1 300 0.8 樟子松林
Pinus sylvestris var.
mongolica forest290 10 22.90 23.18 7樟2落1白 716 0.6 
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表 2 土壤理化性质
Table 2. Physical and chemical properties of Soil
树种类型
Tree types月份
Month温度
temperature/℃含水率
water
content/%酸碱度
pH全碳
TOC/
（mg·g−1）全氮
TN/（mg·g−1）全磷
TP/（mg·g−1）铵态氮
NH+ 4-N/
（mg·kg−1）硝态氮
NO− 3-N/
（mg·kg−1）有效磷
AP/
（mg·kg−1）兴安落叶松林
Larix gmelinii
forest5 4.44 ± 0.15 45.86 ± 0.16 4.23 ± 0.25 31.71 ± 8.64 3.27 ± 1.05 0.73 ± 0.34 17.40 ± 0.61 1.92 ± 0.82 60.48 ± 33.05 6 10.67 ± 0.15 45.69 ± 0.13 4.27 ± 0.17 25.53 ± 8.5 3.09 ± 0.73 0.86 ± 0.5 12.45 ± 0.58 276 ± 0.24 42.49 ± 6.39 7 15.43 ± 0.01 40.81 ± 0.11 4.25 ± 0.15 29.03 ± 17.51 2.37 ± 0.96 0.54 ± 0.22 9.97 ± 0.43 2.63 ± 0.11 35.20 ± 13.76 8 13.43 ± 0.01 35.62 ± 0.15 4.27 ± 0.1 44.28 ± 10.81 3.69 ± 1.13 0.93 ± 0.44 24.91 ± 0.93 1.90 ± 0.70 34.61 ± 8.48 9 6.83 ± 0.17 30.57 ± 0.18 4.37 ± 0.14 37.14 ± 7.66 4.06 ± 1.23 0.89 ± 0.19 23.13 ± 0.54 1.64 ± 0.32 42.12 ± 10.42 樟子松林
Pinus sylvestris
var. mongolica
forest5 4.44 ± 0.17 13.85 ± 0.21 4.41 ± 0.35 30.24 ± 8.06 1.20 ± 0.34 0.33 ± 0.15 10.88 ± 0.31 1.67 ± 0.15 45.63 ± 16.78 6 10.67 ± 0.28 11.53 ± 0.34 4.31 ± 0.25 23.68 ± 2.76 1.19 ± 0.16 0.62 ± 0.31 9.03 ± 0.11 2.54 ± 0.14 40.76 ± 7.78 7 15.43 ± 0.23 9.84 ± 0.16 4.09 ± 0.24 29.54 ± 11.71 1.29 ± 0.49 0.3 ± 0.1 12.31 ± 0.39 2.43 ± 0.15 44.15 ± 25.39 8 13.43 ± 0.23 6.89 ± 0.14 4.27 ± 0.29 38.78 ± 18.32 1.49 ± 0.72 0.74 ± 0.22 12.37 ± 0.57 2.42 ± 0.15 51.30 ± 17.44 9 6.83 ± 0.02 6.39 ± 0.17 4.67 ± 0.18 31.61 ± 9.72 1.52 ± 0.27 0.3 ± 0.05 13.60 ± 0.35 1.76 ± 0.21 33.76 ± 12.46
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表 3 C、N、P含量及其生态化学计量相关系数
Table 3. C, N, P contents and their ecological stoichiometric correlation coefficients between
树种 Tree species 叶 leaf C N P C∶N C∶P N∶P 兴安落叶松 Larix gmelinii C 1 −0.547** −0.456** 0.503** 0.460** −0.274* N 1 0.837** −0.885** −0.806** 0.428** P 1 −0.579** −0.953** −0.122 C∶N 1 0.632** −0.676** C∶P 1 .115 N∶P 1 樟子松 Pinus sylvestris var. mongolica C 1 −0.024 −0.658** 0.209 0.726** 0.619** N 1 0.125 −0.964** −0.124 0.360** P 1 −0.244 −0.952** −0.846** C∶N 1 0.234 −0.249 C∶P 1 0.874** N∶P 1 注：**表示在 0.01 水平（双侧）上显著相关，*表示在 0.05 水平（双侧）上显著相关。下同。Notes: **, means significantly correlated at 0.01 level (bilateral), * means significantly correlated at 0.05 level (bilateral). The same below.
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表 4 C、N、P生态化学计量与土壤因子相关性分析
Table 4. C, N, P ecological stoichiometry and environmental factors correlation analysis between???
树种 Tree species 指标 Indicator 含水量 SWC 温度 ST 铵态氮 NH4 +－N 硝态氮 NO3 −－N 兴安落叶松 Larix gmelinii C −0.765** 0.122 0.416** −0.295* N 0.729** −0.169 −0.199 0.147 P 0.436** −0.544** −0.01 −0.099 C∶N −0.744** −0.181 0.246 −0.258* C∶P −0.374** 0.400** 0.039 0.114 N∶P 0.651** 0.575** −0.318* 0.440** 樟子松 Pinus sylvestris var. mongolica C −0.541** 0.296* 0.182 −0.071 N −0.439** 0.187 0.427** −0.124 P 0.479** −0.079 −0.159 0.159 C∶N 0.354** −0.071 −0.335** 0.087 C∶P −0.659** 0.142 0.263* −0.243 N∶P −0.827** 0.204 0.467** −0.307*
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