典型阔叶红松林主要树种叶性状的垂直变异及经济策略

doi:10.17521/cjpe.2019.0307

摘要/Abstract

摘要：

分析不同树种叶片性状的变化有助于了解植物群落结构。该文通过对典型阔叶红松(Pinus koraiensis)林15种阔叶树种的比叶质量、叶片厚度、叶干物质含量、叶绿素含量指数、叶片碳、氮、磷含量的测定, 分析了冠层高度对叶性状及叶性状间相关关系的影响。结果表明, 水曲柳(Fraxinus mandshurica)和大青杨(Populus ussuriensis)上层的比叶质量显著大于下层, 而其他树种冠层间的比叶质量无显著变化; 叶绿素含量指数在白桦(Betula platyphylla)和春榆(Ulmus japonica)冠层间的分布分别为上层显著大于下层和上层显著大于中层; 单位质量氮含量在水曲柳的中层显著大于上层。叶片性状间存在着广泛的相关性, 比叶质量与叶片厚度、干物质含量在三层间均呈显著正相关关系, 而有些性状, 只在一或二个冠层中存在一定的相关性。山杨(Populus davidiana)和大青杨的叶片倾向于选择光合能力较低、营养浓度较低、呼吸速率较慢的一端, 而黄檗(Phellodendron amurense)和山槐(Maackia amurensis)叶片更倾向于光合能力强、营养物质浓度高的一端。不同树种对光照响应的差异可能会改变不同冠层中叶片的形态和化学性状, 从而有助于群落构建和物种共存。

关键词: 比叶质量, 叶绿素含量, 养分含量, 冠层, 叶经济型谱

Abstract:

Aims An analysis of the variations in leaf traits of different tree species contributes to the understanding of plant community structures.
Methods This study explored the effect of the light environment on leaf traits at different canopy heights in a typical mixed broadleaved-Korean pine forest. We measured the leaf mass per area (LMA), leaf thickness (LT), leaf dry matter content (LDMC), chlorophyll content (CCI), leaf carbon content (C), mass-based and area-based nitrogen content (N_mass and N_area), and mass-based and area-based phosphorus content (P_massand P_area) for 15 broad-leaved tree species in this forest.
Important findings We found that the LMA of Fraxinus mandschurica and Populus ussuriensis in the upper canopy were significantly higher than that in the lower canopy, but no significant vertical changes were detected in other species within the canopy. The CCI of Betula platyphylla increased from the lower to upper canopy. In contrast, the CCI of Ulmus japonica in the upper canopy was significantly higher than that in the middle canopy. The N_mass of Fraxinus mandschurica in the middle canopy was significantly higher than that in the upper canopy. These results indicated that the variations in leaf traits within the canopy were different among species. The LMA was positively correlated with the LT and LDMC in all three canopies, but the correlations of other trait combinations were only significant in one or two canopies. The results suggest that the leaves of Populus davidiana and Populus ussuriensis tend to adopt survival strategies involving lower photosynthetic ability, nutrient concentrations, and respiratory rates, while the leaves of Phellodendron amurense and Maackia amurensis tend to be located at the other end of the economics spectrum with contrasting trait values. Differences in species response to light may alter leaf morphological and chemical traits in canopy layers, and hence contribute to community assembly and species coexistence.

Key words: leaf mass per area, chlorophyll content, nutrient content, canopy, leaf economics spectrum

荀彦涵, 邸雪颖, 金光泽. 典型阔叶红松林主要树种叶性状的垂直变异及经济策略. 植物生态学报, 2020, 44(7): 730-741. DOI: 10.17521/cjpe.2019.0307

XUN Yan-Han, DI Xue-Ying, JIN Guang-Ze. Vertical variation and economic strategy of leaf trait of major tree species in a typical mixed broadleaved-Korean pine forest. Chinese Journal of Plant Ecology, 2020, 44(7): 730-741. DOI: 10.17521/cjpe.2019.0307

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2019.0307

https://www.plant-ecology.com/CN/Y2020/V44/I7/730

图/表 7

表1 典型阔叶红松(Pinus koraiensis)林主要阔叶树种胸径和树高的统计学信息

Table 1 Statistical information of DBH and height for major broadleaf tree species in a typical mixed broadleaved-Korean pine forest

树种 Species	缩写 Abbreviation	胸径 DBH (cm)	树高 Tree height (m)	重要值 Important value (%)
白桦 Betula platyphylla	BP	21.2 ± 3.7	16.6 ± 1.1	0.09
风桦 Betula costata	BC	25.0 ± 3.9	17.7 ± 0.9	3.73
春榆 Ulmus japonica	UJ	48.0 ± 26.7	20.9 ± 4.2	1.42
水曲柳 Fraxinus mandschurica	FM	45.3 ± 14.2	21.2 ± 2.6	2.61
蒙古栎 Quercus mongolica	QM	38.3 ± 8.0	20.3 ± 1.5	0.33
山杨 Populus davidiana	PD	27.5 ± 3.8	18.3 ± 0.9	0.19
大青杨 Populus ussuriensis	PU	28.6 ± 7.4	18.4 ± 1.6	0.97
糠椴 Tilia mandshurica	TM	29.3 ± 4.5	18.7 ± 1.1	0.85
紫椴 Tilia amurensis	TA	43.3 ± 22.2	20.6 ± 3.0	5.13
胡桃楸 Juglans mandshurica	JM	30.2 ± 5.3	18.9 ± 2.2	0.03
山槐 Maackia amurensis	MA	12.2 ± 3.0	13.0 ± 1.7	0.26
黄檗 Phellodendron amurense	PA	25.6 ± 6.5	17.7 ± 1.7	0.21
裂叶榆 Ulmus laciniata	UL	52.1 ± 12.2	22.3 ± 1.4	4.65
花楸 Sorbus pohuashanensis	SP	22.6 ± 8.6	16.5 ± 3.3	0.50
五角槭 Acer pictum subsp. mono	AM	22.2 ± 7.5	16.7 ± 2.2	8.25

表2 典型阔叶红松(Pinus koraiensis)林主要阔叶树种的叶片性状特征

Table 2 Leaf traits for major broadleaf tree species in a typical mixed broadleaved-Korean pine forest

叶性状 Leaf trait	缩写 Abbreviation	单位 Unit	平均值 Mean	标准误差 SE	最大值 Max	最小值 Min
比叶质量 Leaf mass per area	LMA	kg·m^-2	0.049	0.014	0.140	0.016
叶片厚度 Leaf thickness	LT	mm	0.126	0.038	0.298	0.034
叶干物质含量 Leaf dry mass content	LDMC	g·g^-1	0.286	0.056	0.509	0.109
叶绿素含量指数 Chlorophyll content index	CCI		16.83	4.364	34.42	3.14
叶碳含量 Leaf carbon content	C	mg·g^-2	475.6	32.15	554.9	384.3
单位质量的叶氮含量 Leaf nitrogen content per leaf mass	N_mass	mg·g^-2	28.16	6.405	48.43	15.00
单位面积的叶氮含量 Leaf nitrogen content per leaf area	N_area	mg·cm^-2	0.134	0.043	0.385	0.042
单位质量的叶磷含量 Leaf phosphorus content per leaf mass	P_mass	mg·g^-2	1.986	0.699	6.314	0.391
单位面积的叶磷含量 Leaf phosphorus content per leaf area	P_area	mg·cm^-2	0.009	0.004	0.032	0.001

图1 典型阔叶红松(Pinus koraiensis)林不同树种冠层内的叶性状值(平均值+标准误差)。小写字母代表同一树种不同冠层之间叶性状的差异显著性(α = 0.05)。 C, 叶碳含量; CCI, 叶绿素含量指数; LDMC, 叶干物质含量; LMA, 比叶质量; LT, 叶片厚度; Narea, 单位面积的叶氮含量; Nmass, 单位质量的叶氮含量; Parea, 单位面积的叶磷含量; Pmass, 单位质量的叶磷含量。横坐标物种名称缩写见表1。

Fig. 1 Difference of leaf traits (mean + SE) across the canopy layers for different tree species in a typical mixed broadleaved- Korean pine forest. The lowercase letters indicate whether the trait differences were significant across the canopy layers (α = 0.05). C, leaf carbon content; CCI, chlorophyll content index; LDMC, leaf dry mass content; LMA, leaf mass per area; LT, leaf thickness; Narea, leaf nitrogen content per leaf area; Nmass, leaf nitrogen content per leaf mass; Parea, leaf phosphorus content per leaf area; Pmass, leaf phosphorus content per leaf mass. The full names and abbreviations of species on the horizontal axis are shown in Table 1.

图2 典型阔叶红松(Pinus koraiensis)林主要阔叶树种叶片结构性状与叶绿素含量指数的关系。 CCI, 叶绿素含量指数; LDMC, 叶干物质含量; LMA, 比叶质量; LT, 叶片厚度。

Fig. 2 Relationships among leaf structural traits and chlorophyll content index (CCI) for major broadleaf tree species in a typical mixed broadleaved-Korean pine forest. CCI, chlorophyll content index; LDMC, leaf dry mass content; LMA, leaf mass per area; LT, leaf thickness.

图3 典型阔叶红松(Pinus koraiensis)林主要阔叶树种叶片结构性状、叶绿素含量指数与化学性状的关系。 C, 叶碳含量; CCI, 叶绿素含量指数; LDMC, 叶干物质含量; LMA, 比叶质量; LT, 叶片厚度; Narea, 单位面积的叶氮含量; Nmass, 单位质量的叶氮含量; Parea, 单位面积的叶磷含量; Pmass, 单位质量的叶磷含量。

Fig. 3 Relationships among leaf structural traits, chlorophyll content index (CCI) and nutrient contents for major broadleaf tree species in a typical mixed broadleaved-Korean pine forest. C, leaf carbon content; CCI, chlorophyll content index; LDMC, leaf dry mass content; LMA, leaf mass per area; LT, leaf thickness; Narea, leaf nitrogen content per leaf area; Nmass, leaf nitrogen content per leaf mass; Parea, leaf phosphorus content per leaf area; Pmass, leaf phosphorus content per leaf mass.

图4 典型阔叶红松(Pinus koraiensis)林主要阔叶树种叶片化学性状的关系。 C, 叶碳含量; Narea, 单位面积的叶氮含量; Nmass, 单位质量的叶氮含量; Parea, 单位面积的叶磷含量; Pmass, 单位质量的叶磷含量。

Fig. 4 Relationships among leaf chemical traits for major broad-leaved tree species in a typical mixed broadleaved-Korean pine forest. C, leaf carbon content; Narea, leaf nitrogen content per leaf area; Nmass, leaf nitrogen content per leaf mass; Parea, leaf phosphorus content per leaf area; Pmass, leaf phosphorus content per leaf mass.

图5 典型阔叶红松(Pinus koraiensis)林主要阔叶树种叶片性状的主成分分析。 C, 叶碳含量; CCI, 叶绿素含量指数; LDMC, 叶干物质含量; LMA, 比叶质量; LT, 叶片厚度; Narea, 单位面积的叶氮含量; Nmass, 单位质量的叶氮含量; Parea, 单位面积的叶磷含量; Pmass, 单位质量的叶磷含量。B中物种名称缩写见表1。

Fig. 5 Principal component analysis of leaf traits for major broadleaf tree species in a typical mixed broadleaved-Korean pine forest. C, leaf carbon content; CCI, chlorophyll content index; LDMC, leaf dry mass content; LMA, leaf mass per area; LT, leaf thickness; Narea, leaf nitrogen content per leaf area; Nmass, leaf nitrogen content per leaf mass; Parea, leaf phosphorus content per leaf area; Pmass, leaf phosphorus content per leaf mass. The full names and abbreviations of species in panel B are shown in Table 1.

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