植株大小、枝龄和环境共同驱动红松枝性状的变异

doi:10.17521/cjpe.2020.0173

摘要/Abstract

摘要：

许多枝性状的变异受植株大小、枝龄或环境的影响, 但少有研究同时评估这些因素对枝性状种内变异的重要性。该研究以红松(Pinus koraiensis)为研究对象, 通过测定69株胸径(DBH) 0.3-100.0 cm范围内植株不同年龄枝的形态性状、化学性状和解剖性状, 探讨植株大小(DBH或树高)、枝龄与环境因素(光照强度、土壤养分及土壤含水率)对枝性状的影响。结果表明: (1) DBH与树高对枝性状的影响存在差异: 木质密度(WD)、木质部面积占比(R_XA)、韧皮部面积占比(R_PHA)及髓面积占比(R_PA)对DBH更敏感, 而树脂道总面积占比(R_RC)和枝氮含量(WN)受树高影响更大; (2)枝龄是导致红松枝性状种内变异的最主要因素, 植株大小次之, 而环境因素的影响最小; (3) WD、R_PHA与DBH显著正相关, R_PA与DBH显著负相关, R_RC、WN与树高显著正相关; 除WN外, 其余枝性状与枝龄均显著相关, 且随着树木生长, R_RC随枝龄增大而减小的速率加剧, 相反, R_PA随枝龄增大而减小的速率减缓。研究结果有助于了解局域尺度上枝性状种内变异的影响因素以及枝条应对环境变异的适应机制。

关键词: 枝龄, 胸径, 树高, 环境, 木质密度, 解剖性状, 枝氮含量

Abstract:

Aims Variations of many branch traits are affected by plant size, branch age and environment factors, but the relative importance of these factors to intraspecies variations of branch traits has rarely been evaluated simultaneously.
Methods In this study, we took Pinus koraiensis as the research object, to explore the effects of plant size (diameter at breast height (DBH) or tree height), branch age and environmental factors (light intensity, soil nutrient content and water availability) on branch traits, by measuring morphological traits, chemical traits and anatomical traits in different branch ages of 69 individuals with DBH in the range of 0.3-100.0 cm.
Important findings Our results showed that: (1) DBH and tree height had different effects on branch traits: wood density (WD), the xylem area-to-total cross-sectional area ratio (R_XA), the phloem area-to-total cross-sectional area ratio (R_PHA) and the pith area-to-total cross-sectional area ratio (R_PA) were more sensitive to DBH, while the total resin canal area-to-total cross-sectional area ratio (R_RC) and wood nitrogen content (WN) were more affected by the tree height; (2) branch age was the most important factor in driving intra-specific variations of branch traits of P. koraiensis, followed by plant size, while the impact of environment factors was minimal; (3) WD and R_PHA were significantly positively correlated with DBH, while R_PA was significantly negatively correlated with DBH; and R_RC and WN were significantly positively correlated with tree height. Except for WN, the relationships between branch traits and branch age were significant, and as tree growth, the rate of R_RC decreasing with branch age was enhanced, but the rate of R_PA decreasing with branch age was weakened. The results of our study are helpful to understand the driving factors of intraspecific variation of branch traits at the local scale and the adaptation mechanism of branches to cope with environmental changes.

Key words: branch age, diameter at breast height, tree height, environment, wood density, anatomical trait, wood nitrogen content

于青含, 金光泽, 刘志理. 植株大小、枝龄和环境共同驱动红松枝性状的变异. 植物生态学报, 2020, 44(9): 939-950. DOI: 10.17521/cjpe.2020.0173

YU Qing-Han, JIN Guang-Ze, LIU Zhi-Li. Plant size, branch age and environment factors co-drive variations of branch traits of Pinus koraiensis. Chinese Journal of Plant Ecology, 2020, 44(9): 939-950. DOI: 10.17521/cjpe.2020.0173

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https://www.plant-ecology.com/CN/Y2020/V44/I9/939

图/表 7

图1 光学显微镜下不同龄级红松枝解剖结构示意图。A, 当年生枝。B, 二年生枝。C, 三年生枝。D, 四年生枝。

Fig. 1 Schematic diagram of anatomical structure of branches of different ages under an optical microscope for Pinus koraiensis. A, Current-year branch. B, Two-years-old branch. C, Three-years-old branch. D, Four-years-old branch.

表1 红松胸径(DBH)或树高与枝性状回归分析的赤池信息量准则(AIC)值

Table 1 Akaike information criterion (AIC) values in the regressions of diameter at breast height (DBH) or tree height against each branch traits of Pinus koraiensis

性状 Trait	DBH (cm)	树高 Tree height (m)
木质密度 WD (g·cm^-3)	-1 025	-1 022
枝氮含量 WN (mg·g^-1)	1 282	1 272
木质部面积占比 R_XA	-327	-325
韧皮部面积占比 R_PHA	-1 090	-1 086
髓面积占比 R_PA	-1 364	-1 362
树脂道总面积占比 R_RC	-1 129	-1 142

表2 木质密度(WD)、枝氮含量(WN)、木质部面积占比(RXA)、韧皮部面积占比(RPHA)、髓面积占比(RPA)以及树脂道总面积占比(RRC)的统计信息

Table 2 Statistical information of wood density (WD), wood nitrogen content (WN), the xylem area-to-total cross-sectional area ratio (RXA), the phloem area-to-total cross-sectional area ratio (RPHA), the pith area-to-total cross-sectional area ratio (RPA) and the total resin canal area-to-total cross-sectional area ratio (RRC)

性状 Trait	最大值 Maximum	最小值 Minimum	平均值 Mean (标准偏差 SD)	变异系数 Coefficient of variation (%)
WD (g·cm^-3)	0.548	0.272	0.39 (0.03)	8
WN (mg·g^-1)	19.080	1.067	8.99 (3.72)	41
R_XA	0.634	0.092	0.31 (0.12)	39
R_PHA	0.146	0.026	0.09 (0.02)	22
R_PA	0.098	0.005	0.03 (0.01)	33
R_RC	0.136	0.000	0.07 (0.02)	29

图2 红松当年生和多年生枝性状随植株大小(胸径或树高)的变异。*, p > 0.05; **, p > 0.01; ***, p > 0.001。

Fig. 2 Variations of branch traits in current-year and old branches with plant size (DBH or tree height) for Pinus koraiensis. DBH, diameter at breast height; RPA, pith area-to-total cross-sectional area ratio; RPHA, phloem area-to-total cross-sectional area ratio; RRC, total resin canal area-to-total cross-sectional area ratio; RXA, xylem area-to-total cross-sectional area ratio; WD, wood density; WN, wood nitrogen content. *, p > 0.05; **, p > 0.01; ***, p > 0.001.

表3 红松枝性状与植株大小、枝龄和环境因素(光照强度、土壤含水量、土壤氮含量和磷含量)之间的广义线性模型(GLM)

Table 3 Generalized linear models (GLM) among branch traits, tree size, branch age and environment factors (light availability, soil water content, soil nitrogen content, soil phosphorus content) for Pinus koraiensis

性状 Trait	胸径 DBH (cm)	树高 Tree height (m)	枝龄 Branch age (year)	光照强度 Light intensity (mol·m^-2·d^-1)	土壤含水量 Soil water content (g·g^-1)	土壤氮含量 Soil nitrogen content (mg·g^-1)	土壤磷含量 Soil phosphorus content (mg·g^-1)	截距 Intercept
木质密度 WD (g·cm^-3)	0.007^**		0.009^***	0.003	0.003	-0.001	-0.001	0.393^***
枝氮含量 WN (mg·g^-1)		1.732^***	-0.373	0.121	-0.062	0.302	-0.224	8.881^***
木质部面积占比 R_XA	-0.006		0.090^***	-0.009	0.005	0.020^**	-0.019^*	0.310^***
韧皮部面积占比 R_PHA	0.006^***		0.007^***	-0.005^**	0.005^*	-0.001	-0.003	0.087^***
髓面积占比 R_PA	-0.003^***		-0.008^***	>0.001	>0.001	-0.001	>0.001	0.033^***
树脂道总面积占比 R_RC		0.011^***	-0.008^***	0.002	>0.003	-0.003	-0.001	0.067^***

图3 不同植株大小(胸径(DBH)或树高)红松枝性状随枝龄的变异。*, p > 0.05; **, p > 0.01; ***, p > 0.001。

Fig. 3 Variations of branch traits of different plant sizes (DBH or tree height) with branch age for Pinus koraiensis. DBH, diameter at breast height; RPA, pith area-to-total cross-sectional area ratio; RPHA, phloem area-to-total cross-sectional area ratio; RRC, total resin canal area-to-total cross-sectional area ratio; RXA, xylem area-to-total cross-sectional area ratio; WD, wood density; WN, wood nitrogen content. *, p > 0.05; **, p > 0.01; ***, p > 0.001.

图4 红松枝性状随枝龄的变化斜率随不同植株大小(DBH或树高)的变异趋势。RPA, 髓面积占比; RPHA, 韧皮部面积占比; RRC, 树脂道总面积占比; RXA, 木质部面积占比; WD, 木质密度。*, p > 0.05; **, p > 0.01。

Fig. 4 Variation trend of the slope of branch traits against branch age with different plant sizes (DBH or tree height) for Pinus koraiensis. DBH, diameter at breast height; RPA, pith area-to-total cross-sectional area ratio; RPHA, phloem area-to-total cross-sectional area ratio; RRC, total resin canal area-to-total cross-sectional area ratio; RXA, xylem area-to-total cross-sectional area ratio; WD, wood density. *, p > 0.05; **, p > 0.01.

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