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近日,北京林业大学林学院田地青年研究员及合作者在Nature Communications上发表Article论文“Environmental versus phylogenetic controls on leaf nitrogen and phosphorous concentrations in vascular plants”。田地为论文第一作者兼第一通讯作者,北京林业大学(林木资源高效生产全国重点实验室)为第一完成单位。
氮和磷是植物细胞基本结构的重要组分,二者的平衡关系影响植物个体的生长发育、土壤–植物间的养分循环和生态系统碳氮磷循环等重要过程。叶片氮和磷浓度及其计量关系是表征植物功能的关键性状。得益于大量野外调查数据,前人报道了不同研究尺度植物叶片氮磷计量特征在环境梯度上的变化规律。然而,其驱动机制尚未得到解析,尤其是区分并量化环境和系统发育对维管植物叶片氮磷计量特征的驱动力,依然是植物生态化学计量学(田地、严正兵、方精云 综述:植物生态化学计量特征及其主要假说)中的难题。
Fig. 1: Two hypotheses of leaf N and P stoichiometry variations expressed in a trait gradient analysis. a, b Indicate the Biogeochemical Niche Hypothesis and Stoichiometric plasticity, respectively. Each colored line represents species-specific linear regressions of a species’ leaf N (P or N:P) vs the mean leaf N (P or N:P) across all species occurring at the respective site.
在前期全球植物叶片氮磷数据库的基础上(Tian et al. 2019. Ecology),联合欧洲ICP调查数据(Sardans et al. 2021. Nature Ecology & Evolution)和植物功能属性数据库(TRY-Plant Trait Dataset),构建并升级完成目前全球最大的植物叶片氮磷含量匹配数据库,包含36,413 组记录,每组记录匹配 45 个环境变量,共覆盖全球 7,549 个研究站点,涉及3,700个物种(全部数据已开源共享:Stocker, B. D., & Tian, D. geco-bern/leafnp_data: v1.0: Initial release (v1.0) [Data set]. Zenodo https://doi.org/10.5281/zenodo. 11071944 (2024))。
Fig S2. Distribution of sites from which leaf N and P data were sourced, plotted in climatic space defined by mean annual temperature (x-axis) and mean annual precipitation (y-axis).
在此基础上,同时采用传统上常用的线性模型(Linear Models,LMs)和新型机器学习中随机森林(Random Forest,RF),对比二者在分解环境因素和物种特性对叶片氮磷计量关系宏观格局的解释率方面的差异。结果发现,叶片氮磷计量特征在物种内的变异与种间变异一样重要,沿环境梯度的种内可塑性因物种而异,总体上氮磷比可塑性最高,而叶氮含量可塑性最低。环境变量分别解释了叶氮、磷浓度及其比值变化的 29%、31% 和 22%,随机森林模型确定了环境变量对种内变异的影响,而广泛使用的线性混合模型很大程度上忽略了这些影响(原因在于线性混合模型行fixed term和random term的确定较大的主观随意性)。另外,全球变化因子,如CO2浓度升高、大气N沉降量增加等,显著改变了维管植物的叶氮磷计量特征,这些改变很可能会通过影响长期尺度上的物种更替来实现,同时也会通过短期内快速改变物种内的性状变异来实现。(全部代码已开源共享:Stocker, B. D., & Tian, D. geco-bern/leafnp: v1.0: Initial release (v1.0). Zenodo https://doi.org/10.5281/zenodo.11071816 (2024))。
该研究表明环境对驱动物种内叶片氮磷计量关系的可塑性具有不可忽略的影响,对生物地球化学生态位假说(Biogeochemical Niche Hypothesis)以及物种分布在塑造叶片氮磷计量全球格局的绝对地位提出了挑战。这一理论突破对于地球系统模型升级中处理碳氮磷循环相关环节具有重要的启示,尤其是植被动态模型在预测全球环境变化与植被响应间的互馈时,应充分考虑物种内和物种间的植物叶功能属性变化。
Fig. 2: Variable selection and effects. a-c Variable selection order determined by the recursive feature elimination for leaf N (a), P (b), and N:P (c). d-f Effect magnitudes of the selected variables, measured by the t-value of the coefficients determined by the LMMs(Tian et al. unpublished).
Fig. 4: Trait gradient analysis. a–c Regression of log-transformed species-specific leaf N, P, and N:P versus the site-mean values (the dashed line shows the 1:1 relationship). d Distribution of slopes of species-specific regressions. e Distribution of species occurrence, quantified as the ranges of site-level mean leaf N, P, and N:P, normalized by their respective overall mean, along which species occurred. f Species variation decomposition (see “Methods” section), measuring the proportion of variations in leaf N, P, and N:P arising within vs across species. g–i Slopes of the species-specific regressions for species with the largest amount of data (N > 50). Gray lines indicate the 95% confidence interval of the slope estimate. Source data are provided as a Source Data file.
瑞士伯尔尼大学Benjamin Stocker教授为本文的共同通讯作者。本研究的合作者包括中国科学院植物研究所严正兵研究员、瑞士苏黎世大学Bernhard Schmid教授、德国马普研究所Jens Kattge研究员和北京大学方精云院士。该研究得到十四五重点研发青年科学家项目(No. 2022YFD2201600)、国家自然科学基金(32271680,31800397,31901086)等项目的资助和支持。田地得到Swiss Government Excellence Scholarship,中国科协第七届青年人才托举工程项目和北京林业大学“5•5工程”创新团队和青年拔尖人才计划的支持。
论文链接:https://www.nature.com/articles/s41467-024-49665-4
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