陆地生态系统是一个由不同组分相互协调构成的有机整体，理解这些组分之间的协同关系是深入认识生态系统运作的基础。然而，量化生态系统各组分之间的关系一直是生态学所面临的一道难题。基于生态策略和性状的概念为揭示这些关系提供了理论基础和研究框架。相较于已经明确的植物性状权衡的生态策略维度及其与生态系统碳循环的联系，生态系统地下部土壤生物生态策略的研究仍显薄弱。作为地球上数量最丰富的后生动物类群，土壤线虫在全球碳循环过程中同样发挥着不可替代的作用，但其关键的性状-策略维度尚未得到充分解析。植物和线虫同为生态系统碳循环的重要调控者，当研究视角从单一类群扩展至跨界类群时，线虫生态策略的知识空白严重制约了我们对生态系统碳循环的全面理解。那么，二者的性状与生态策略之间究竟存在怎样的协同关系，这种协同关系又如何影响生态系统的碳循环过程？本研究通过构建原创性理论框架并结合大尺度野外采样调查，从生态策略角度系统探索了这一科学问题，旨在全面、系统地理解跨界生态策略与生态系统功能之间的关联。

本研究从两个方面展开，首先，在理论研究方面，系统梳理现有研究中的线虫性状，对其进行归纳分类，并在此基础上量化核心性状之间的权衡关系，解析线虫生态策略的关键维度，同时尝试构建其与植物策略维度之间的联系，从而建立原创性理论框架，为阐明跨界生态策略与土壤碳循环的关系提供新视角。其次，在实证研究方面，依托我国南方典型生态系统样带，开展大尺度野外调查采样工作，系统测定植物性状、线虫性状以及土壤碳循环关键指标，明确植物与线虫生态策略的多维协同规律，并探究跨界生态策略与土壤碳循环之间的联系，最终实证支持本研究提出的理论框架。主要结果如下：

（1）基于理论知识和现有研究，将线虫性状归类为形态、生理、生活史和群落四个类别，解决了当前线虫性状描述不一致、数据碎片化的问题，为线虫性状研究提供了标准化性状分类体系。在此基础上，本研究开创性地提出了“线虫经济系谱”理论框架，聚焦于线虫生长、生存和繁殖策略，涵盖五个关键性状。该系谱描述了一个从“快速投资-收益”到“缓慢投资-收益”的连续系谱：在“快速”端，线虫个体通常体型较小，代谢和繁殖速率较高，寿命较短且产卵较小；而在“缓慢”端，线虫个体则表现为体型和卵较大，寿命较长且代谢和繁殖速率较低。进一步地，本研究建立了植物与线虫经济系谱之间的跨界联系，提出了“整合的植物与线虫经济系谱”框架。在该整合系谱的“快速”端，植物和线虫物种协同促进碳和养分的快速循环，而在碳和养分循环较慢的生态系统中，则以“缓慢”的植物和线虫物种为主导。

（2）采用空间替代时间的方法，在云南大围山自然保护区选取了农田弃耕后自然恢复的草地（先锋期）、15年（前期）、30年（中期）和60年以上（后期）的森林，构建完整的自然恢复时间序列。植物和线虫的生态策略在自然恢复过程中呈现一致的变化轨迹，从先锋期以快速资源获取和高代谢活性的小型生物为主，逐渐演替到后期以大型、缓慢生长的生物为主。这一生态策略的转变伴随着土壤有机碳的显著积累，从先锋期的 39.4 g kg⁻¹ 增至后期的 118.2 g kg⁻¹（p < 0.05）。基于植物和线虫性状的协同变化，研究构建了整合的植物与线虫经济系谱，首次为量化植物与线虫生态策略之间的关系提供了直接证据。在本研究的生态系统自然恢复过程中，随机森林模型表明，整合经济系谱上植物和线虫性状的协同变化是土壤有机碳变化的最佳预测因子（p < 0.001），其解释力优于单独使用其中任何一组性状。方差分解结果进一步表明，76%的土壤有机碳变异可由植物与线虫性状的协同效应解释。此外，结构方程模型揭示了整合的植物与线虫经济系谱通过调控微生物性状，如微生物碳利用效率与微生物生物量，进而影响土壤有机碳的变化。

（3）基于中国南方典型生态系统样带，本研究选取了六个典型样点，涵盖草地、灌丛和森林三种植被类型，共设置87个样地，系统分析了自然恢复序列中植物性状、线虫性状和土壤碳循环关键指标。研究首先确定了植物生态策略、线虫生态策略和土壤碳循环的关键维度。植物根系的主要性状-策略维度包括根系经济系谱和根系协作维度，前者反映了根系快-慢权衡策略，即高代谢活性与组织投资之间的权衡，后者体现了根系在自主获取资源与通过与菌根真菌合作获取资源之间的权衡。这一结果与全球尺度上的多维格局相一致，进一步验证了植物生态策略的多维性。线虫的主要性状-策略维度同样包括两个方面，其一是线虫经济系谱，由体型大小和寿命定义，反映了在体型构建成本与生长潜力之间的快-慢连续体，其二是线虫繁殖维度，体现了雄性比例与后代比例之间的权衡。此外，研究还确定了土壤碳循环的两个主要维度，其中一个维度反映了碳流，涵盖从具有高碳相关酶活性和呼吸作用的快速碳流到缓慢碳流；另一个维度代表了碳库，由土壤有机碳含量的高低定义。线性回归模型揭示了植物根系经济系谱与线虫经济系谱之间存在显著的协同关系（p < 0.001），这一协同的快-慢连续体与土壤碳库的变化密切相关（p < 0.001），而繁殖维度与根系协作维度相契合（p < 0.001），这一协同的协作–繁殖维度主要与土壤碳流密切相关（p < 0.001）。研究结果首次将植物、线虫和土壤碳循环纳入到一个统一的框架中，深化了对全球变化背景下生态系统运作的整体性理解。

综上所述，本研究首次将生态策略框架应用于土壤线虫分析，提供了一种具有理论深度和实践意义的全新思路和方法。更重要的是，本研究以中国南方典型生态系统样带为平台，选取了共22个典型的生态系统作为研究地点，结合野外调查采样和室内实验测定，通过可视化和定量化生态系统不同组分在各自维度上的协调关系，系统阐明了植物与线虫生态策略的跨界协同及其与土壤碳库和碳流的关系，涵盖从一维经济系谱到二维性状空间的全方位分析。具体而言，本研究成功构建了一个涵盖了植物、线虫和土壤碳循环的整合性性状空间的概念框架。这一性状空间不仅为生态恢复提供了基于自然的解决方案，还为改进生态系统功能与全球变化的预测模型奠定了理论基础。此外，这一跨类群、跨学科的概念框架有望整合更多生物类群及生态系统功能，并基于性状-策略维度的连续变化进行扩展。然而，未来仍需在全球尺度更多样的生态系统类型中进行验证、完善和拓展。

Terrestrial ecosystems comprise diverse and interconnected components, and their properties and functioning depend on the tight coordination among them. However, quantitative evidence of the intricate interactions among key ecosystem components remains a central goal of ecology. The lens of ecological strategy schemes allows us to understand how organisms function within ecosystems and quantify their relationships with other organisms, thereby conceptualizing their linkages to terrestrial ecosystem functions. Plant ecologists have pioneered this effort, identifying the fundamental dimensionality of trait variation and exploring how this variation drives the dimensionality of ecosystem functions. However, as for many aspects of ecology, far less is known about the world belowground than aboveground. For example, the traits and ecological strategies of soil nematodes – the most abundant animals on Earth and a group that plays a key role in ecosystem functioning – remain less explored. The knowledge gap hinders our ability to explicitly consider multidimensional cross-taxon between plants and nematodes – two key drivers in global carbon cycle – towards more nuanced predictions of ecosystem carbon cycles in the face of ongoing global challenges. This gives rise to a key scientific question: to what extent are the traits and strategies of plants and nematodes coordinated, and what are the implications for the ecosystem carbon cycle? This study addresses this challenge by developing a conceptual framework and conducting large-scale field sampling from an ecological strategies perspective, aiming to understanding the linkages between cross-kingdom ecological strategies and ecosystem functions.

This study systematically summarizes and categorizes nematode traits into distinct clusters, identifying trade-offs between key traits to determine the main dimensions of nematode ecological strategies. It further explores the coordinated linkages between plant and nematode ecological strategies and their trait proxies, developing a conceptual framework for fully understanding cross-kingdom ecological strategies and the role in soil carbon cycling. Additionally, a large-scale field sampling is conducted at 7 sites across southern China. Functionally important traits of plants and nematodes, along with key aspects of soil carbon cycling, are analyzed to assess the coordination between plant and nematode ecological strategies and to evaluate the linkages between plant-nematode coordination and soil carbon cycling, providing empirical evidence for the proposed conceptual framework. The main findings are as follows:

(1) Drawing on theoretical knowledge and existing studies, this study categorizes nematode traits into morphological, physiological, life history, and community clusters. The standardized classification system aims to propose a unified language for nematode trait ecology. We then propose the “nematode economics spectrum (NES)”, focusing on five key traits related to nematode growth, survival, and reproduction. The NES runs from species with potential for fast growth and fast returns on investment into resource acquisition and metabolism, to species that have the potential to invest in reproduction with slow returns. At the fast end are species with a small body size, high rates of metabolism and reproduction, and short lifespans that produce small eggs. Species at the slow end have large body and egg sizes, long lifespans, and low rates of metabolism and reproduction. Furthermore, we bridge the NES with the plant economics spectrum (PES), forming a PES–NES framework in which plant and nematode species at the fast end of the continuum contribute to fast carbon and nutrient cycling, while ecosystems with slow carbon and nutrient cycling are dominated by species at the slow end.

(2) This study system is a space-for-time chronosequence spanning pioneer (grassland), early (15-year forest), mid (30-year forest), and climax (over 60-year forest) stages of natural restoration on ex-arable lands following agricultural abandonment in the Daweishan National Natural Reserve. Covariation in plant and nematode traits provides compelling evidence for synchronous changes in plants and nematodes during natural restoration, ranging from small-bodied organisms characterized by fast resource acquisition and metabolic activities in the pioneer stage to large and slow-growing organisms in the climax stage. Change in plant and nematode traits correspond to enhanced soil organic carbon (SOC) dynamics, which SOC content significantly increasing across stages from 39.4 to 118.2 g C kg^-1 soil (p < 0.05). We identify an integrated fast-slow economics spectrum encompassing plants and nematodes, providing the first empirical evidence for quantifying the relationship between their ecological strategies. The random forest models demonstrate that the integrated economics spectrum is the most important predictors of SOC (p < 0.001), outperforming either the PES or NES alone. Furthermore, variation partitioning analysis reveal that variation in the fast–slow spectra of plants and nematodes (the joint effects) explained 76% of the total variation in SOC. Finally, structural equation modeling reveals that the integrated fast-slow economics spectrum influenced soil organic carbon through its regulation of microbial traits, including microbial carbon use efficiency and microbial biomass.

(3) This study analyse functionally important traits of plants and nematodes, and key aspects of soil carbon cycling from 87 plots spanning different vegetation types ‒ grasslands, shrublands and forests ‒ along a natural restoration chronosequence at 6 sites in China. The study first identified the primary trait-strategy dimensions of plant roots, soil nematodes and soil carbon cycling. First, we identify two key trait-strategy dimensions for plant roots, one being the root economic spectrum (RES), another is the collaboration dimension. The RES represents the “fast–slow” trade-offs between high metabolic activity and high tissue investment, while the collaboration dimension captures the trade-offs between do-it-yourself resource uptake and resource acquisition being primarily delegated to arbuscular mycorrhizal fungal partners. Our results agree well with the previously reported global root economics space, which depicts different trait dimensions of plant ecological strategies for belowground resource acquisition. Next, we identify two key trait-strategy dimensions for nematodes. One, the NES, is defined by body size and lifespan, reflecting a fast‒slow continuum balancing body construction costs against growth potential. The other, the reproduction dimension, captures a trade-off between male proportion and offspring proportion. We then identify two key dimensions to soil carbon cycling. One represents carbon flux, ranging from fast fluxes with high carbon-related enzyme activities and respiration, to slow fluxes. The second represents the carbon pool, defined by the distribution of soil organic carbon. Linear regression models further confirm that variation along the NES is coordinated with the RES (p < 0.001), with their integrated fast–slow continuum is associated with variation in carbon pools (p < 0.001). The reproduction dimension aligns with the root collaboration dimension (p < 0.001), with this collaboration–reproduction dimension is related to carbon fluxes (p < 0.001). Our study goes beyond focusing solely on plants, nematodes, or soil carbon – it integrates these key components of ecosystems into a unified framework for the first time, providing a comprehensive understanding to ecosystem dynamics amid ongoing and future climatic and environmental changes.

Generally, this study is the first attempt to apply ecological strategy schemes to nematodes, providing new perspectives and methods for nematode ecology. Crucially, by visualizing and quantifying the tight coordination between the dimensionality of each of key ecosystem components and ecosystem functions across 22 typical ecosystem in southern China, it expands previous findings by shifting the focus from the linkages between the unidimensional trait spectrum of individual guilds and ecosystem functions to the connections between the bidimensional trait space of cross-kingdom assembly and the primary axes of variation of ecosystem functions. This study successfully integrates key ecosystem components and functions – plants, nematodes and soil carbon cycling – into a unified framework: an integrated trait space that captures the linkages between cross-kingdom ecological strategies and carbon pools and fluxes. The integrated trait space provides nature-based solutions for ecological restoration, and a foundation for improving predictive models of ecosystem function and global change. With its cross-taxon, cross-disciplinary focus, this trait space has the potential to incorporate additional ecosystem components and ecosystem functions, expanding the framework based on continuous variation along trait-strategy dimensions. However, it is still in its infancy and currently offers limited insights. A broader generalization of the presented findings requires further studies with larger sample sizes across multiple sites and diverse climatic conditions.

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