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    油松异龄叶功能性状对氮磷添加的响应

    Responses of functional traits of different-aged leaves of Pinus tabuliformis to nitrogen and phosphorus addition

    • 摘要:
      目的 揭示油松异龄叶片资源分配特征及其功能性状对养分添加的响应机制,以期为阐明森林生态系统中植物对养分变化的适应策略及其生态学意义提供理论支撑。
      方法 在山西太岳山灵空山林场设置氮(N)、磷(P)添加控制试验,系统测定油松不同年龄叶片的结构性状、色素性状及化学性状3大类共8个叶片功能性状,结合土壤养分动态监测,采用Pearson相关检验和回归分析探讨异龄叶多性状间的协同与权衡关系,利用冗余分析来评估显著影响油松叶功能性状的关键土壤环境因子,解析不同年龄叶片功能性状对养分输入的响应规律。
      结果 (1)在结构性状和色素性状方面:低N处理使当年生针叶的比叶面积(SLA)显著增加18.5%,同时使其叶干物质含量(LDMC)显著降低7.96%,即N添加促进了当年生叶资源获取能力的提升;叶绿素(CHL)含量仅在当年生叶中因N添加而显著提升,而二年生CHL含量在低N处理下显著降低了16.0%;相比之下,P 添加和N + P 联合效应对其影响有限。(2)在化学性状方面:低N、低P添加显著提高了二年生针叶的P含量,高P、N + P添加显著抑制了当年生、一年生针叶的P含量,N、P 和N + P这3种养分添加处理均显著增加了当年生叶的碳(C)含量,但均未显著改变叶片N含量,反映出异龄叶化学性状对外源养分输入响应具有明显差异,其适应过程主要通过P分配调整和C积累变化来实现。(3)不同发育阶段叶片表现出明显的功能分化,当年生叶有较低的叶厚(LT)、LDMC,较高的SLA、叶N含量和叶P含量,趋向“快速投资收益型”的积极策略;往年生叶有较高的LT、LDMC,较低的SLA、叶N含量和叶P含量,倾向“缓慢投资收益型”的保守策略,反映出油松异龄叶通过策略分化适应养分环境变化。(4)叶性状之间普遍存在显著相关性,并在异龄叶之间存在着不同的制约关系,表明叶片功能协同与资源分配关系会随叶龄变化而发生调整。(5)导致不同发育阶段下叶功能性状改变的主导土壤因子并不相同,影响油松当年生叶的关键因子是土壤pH,一年生叶的关键因子是土壤全磷(TP)含量,而二年生叶的关键因子是土壤含水量(SWC),体现了叶片在不同发育阶段对土壤环境的敏感程度存在差异。
      结论 N、P添加对油松针叶功能性状有显著影响,且不同年龄的叶片对养分添加的响应存在差异,这为常绿树种不同年龄的叶片适应N、P沉降的机制提供了新的见解,也将为进一步认识植物在养分变化背景下通过性状协同优化资源配置的过程提供理论支持。

       

      Abstract:
      Objective To elucidate the resource allocation patterns of leaves from different age classes of Pinus tabulaeformis and the mechanisms underlying their functional traits’ responses to nutrient addition, with the aim of providing a theoretical foundation for understanding the adaptive strategies of plants to nutrient fluctuations in forest ecosystems and their ecological significance.
      Method In this study, we established a controlled nitrogen (N) and phosphorus (P) addition experiment in the lingkongshan Forest of Taiyue Mountain, Shanxi Province. We systematically measured eight functional traits across three major categories—structural, pigment, and chemical characteristics—in leaves of different ages from Pinus tabulaeformis. Combining this with dynamic monitoring of soil nutrients, we employed Pearson correlation tests and regression analysis to explore the synergistic and trade-off relationships among multiple traits in leaves of different ages. Additionally, we utilized redundancy analysis to identify key soil environmental factors significantly influencing the functional traits of Pinus tabulaeformis leaves and to elucidate the response patterns of these traits to nutrient inputs.
      Result (1) Regarding structural and pigment characteristics, low N treatment significantly increased the specific leaf area (SLA) of current-year needles by 18.5%, while simultaneously reducing their leaf dry matter content (LDMC) by 7.96%; indicating that nitrogen application enhanced the resource acquisition capacity of current-year leaves; Chlorophyll (CHL) content increased significantly only in current-year leaves with N addition, while second-year CHL decreased significantly by 16.0% under low N treatment. In contrast, P addition and the combined N + P effect had limited impact. (2) Regarding chemical properties, low N and low P addition significantly increased P content in second-year needles. High P and N + P additions significantly suppressed P content in both current-year and one-year-old needles. All three nutrient addition treatments significantly increased carbon (C) content in current-year leaves, but none significantly altered leaf N content. This reflects distinct differences in the response of leaves of different ages to exogenous nutrient inputs, with the adaptation process primarily achieved through adjustments in P allocation and changes in carbon accumulation. (3) Leaves at different developmental stages showed marked functional differentiation. Current-year leaves had lower leaf thickness (LT) and LDMC, but higher SLA, leaf N content, and leaf P content, indicating an acquisitive “fast investment-return” strategy. In contrast, older leaves had higher LT and LDMC, but lower SLA, leaf N content, and leaf P content, reflecting a conservative “slow investment-return” strategy. These findings indicate that leaves of different ages in Pinus tabulaeformis adapt to changes in the nutrient environment through strategic differentiation. (4) Leaf traits were generally significantly correlated, and the trade-off relationships among traits varied with leaf age, indicating that functional coordination and resource allocation patterns shift across developmental stages. (5) The dominant soil factors driving changes in leaf functional traits differed among leaf ages: soil pH was the main factor for current-year leaves, soil total phosphorus (TP) for 1-year-old leaves, and soil water content (SWC) for 2-year-old leaves. These results suggest that leaves at different developmental stages differ in their sensitivity to soil environmental conditions.
      Conclusion Nitrogen and phosphorus additions significantly affected the functional traits of Pinus tabuliformis needles, and leaves of different ages showed distinct responses to nutrient inputs. These findings provide new insights into the mechanisms by which evergreen species with leaves of different ages adapt to nitrogen and phosphorus deposition, and offer a theoretical basis for further understanding how plants optimize resource allocation through trait coordination under changing nutrient conditions.

       

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