长白落叶松−水曲柳混栽对长白落叶松生长发育及防御能力的影响

Effects of Larix olgensis-Fraxinus mandshurica mixed planting on the growth, development and defense capacity of Larix olgensis

摘要

【目的】探究长白落叶松−水曲柳混栽对长白落叶松生长发育及防御能力的影响，揭示混栽环境下长白落叶松生长特征与防御策略的动态变化规律，以期为东北地区长白落叶松人工林的混交配置、可持续经营及生态防护林建设提供科学理论依据与实践参考。【方法】本试验以盆栽幼树模拟混栽样地，设置长白落叶松−水曲柳 1∶1 混栽（HZ1）、3∶3 混栽（HZ3）及长白落叶松对照处理，测定长白落叶松的生长发育指标（苗高、地径、各器官干质量及生物量分配格局）、叶片营养物质含量（氨基酸、可溶性糖、蛋白质）、物理防御指标（比叶重、叶片厚度、叶片密度）以及化学防御指标（抗氧化酶活性、次生物质含量），系统探究不同混栽处理对长白落叶松生长发育与防御特征的影响差异。【结果】（1）与对照组相比，混栽后长白落叶松的树高及各器官干质量显著升高（ P < 0.05），但地径显著降低（ P < 0.05）；生物量向叶片和根系倾斜分配，体现了混生环境下植株优先争夺光照与养分的适应性策略；同时针叶比叶重和叶片厚度显著提高（ P < 0.05），物理防御屏障得到强化。（2）混栽后长白落叶松针叶内氨基酸、可溶性糖及蛋白质含量整体高于对照组，表明混栽提升了植株的生理代谢活性与营养储备水平。（3）长白落叶松针叶内抗虫性次生物质（单宁、木质素）含量高于对照组，其中单宁含量在多数时间点显著高于对照组（ P < 0.05）；而与抗逆、抗氧化相关的总酚含量显著低于对照组（ P < 0.05），抗虫化学防御能力增强；抗氧化酶（CAT、POD、SOD）的活性整体低于对照组，反映混交林环境胁迫压力更小，植株实现了资源的优化配置。【结论】长白落叶松−水曲柳混栽可协同提升长白落叶松的生长能力与防御能力，植株通过调整生物量分配与防御物质合成策略，实现了生长与防御的优化平衡。研究结果证实了针阔混交模式在提升人工林生产力与抗逆性方面的双重优势，可为东北林区长白落叶松人工林的结构优化与可持续经营提供了理论依据与实践指导。

Abstract

[Objective] To investigate the effects of mixed planting of Larix olgensis and Fraxinus mandshurica on the growth, development and defense capacity of the former, and to reveal the dynamic patterns of growth characteristics and defense strategies of L. olgensis under the mixed planting conditions, with the aim of providing scientific theoretical basis and practical references for mixed planting configuration, sustainable management and ecological shelterbelt construction of L. olgensis plantations in Northeast China. [Method] Potted saplings were used to simulate mixed planting plots. Three treatments were set: 1∶1 mixed planting (HZ1) and 3∶3 mixed planting (HZ3) of L. olgensis and F. mandshurica, and monoculture control. Growth and development indices (including seedling height, ground diameter, dry weight of various organs and biomass allocation pattern), nutrient contents in leaves (amino acids, soluble sugars, proteins), physical defense indices (leaf mass per area, leaf thickness, leaf density), and chemical defense indices (antioxidant enzyme activity, secondary metabolite content) of L. olgensis were determined. The differences in growth, development, and defense-related characteristics of L. olgensis under different mixed planting patterns were systematically investigated. [Result] (1) Compared with the control group, L. olgensis in mixed planting showed significantly greater tree height and dry mass of all organs (P < 0.05), but a significantly smaller ground diameter (P < 0.05). Biomass allocation was biased toward leaves and roots, reflecting the adaptive strategy of plants to prioritize competition for light and nutrients in mixed environments. Meanwhile, leaf mass per area and leaf thickness of needles were significantly higher in the mixed planting treatment (P < 0.05), indicating the enhancement of physical defense barriers. (2) The contents of amino acids, soluble sugars, and proteins in the needles of L. olgensis under mixed planting were generally higher than those in the control group, indicating that mixed planting enhanced the physiological metabolic activity and nutrient reserve levels of the plants. (3) The contents of insect-resistance secondary metabolites (tannins and lignin) in the needles of L. olgensis were higher than those in the control group, with tannin content being significantly higher at most time points (P < 0.05). In contrast, the total phenol content associated with stress resistance and antioxidant activity was significantly lower (P < 0.05), indicating enhanced chemical defense capability against insects. The activities of antioxidant enzymes (CAT, POD, and SOD) were generally lower than those in the control group, reflecting smaller environmental stress pressure in mixed forests and optimized resource allocation by the plants. [Conclusion] Mixed planting of L. olgensis and F. mandshurica can synergistically enhance the growth capacity and defense capacity of L. olgensis. The plants achieved an optimized balance between growth and defense by adjusting biomass allocation and defense substance synthesis strategies. The research results verified the dual advantages of coniferous-broadleaved mixed planting patterns in enhancing plantation productivity and stress resistance from the perspective of physiological and ecological mechanisms, providing theoretical basis and practical guidance for structural optimization and sustainable management of L. olgensis plantations in Northeast forest regions.

关键词

长白落叶松 / 水曲柳 / 混栽 / 生长发育 / 物理防御 / 化学防御

Key words

Larix olgensis / Fraxinus mandshurica / mixed planting / growth and development / physical defense / chemical defense

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基金资助

黑龙江省自然科学基金联合基金培育项目(PL2024D001)