叶片病原真菌对植物物种共存的影响: 进展与挑战

doi:10.17520/biods.2022525

摘要/Abstract

摘要：

群落内物种如何共存是群落生态学研究中最具争议的核心问题之一。根据当代物种共存理论框架, 维持物种共存的机制可以分为稳定化和均等化机制。尽管植物叶片病原真菌在自然界中大量存在, 但是目前尚不完全清楚叶片病原真菌如何通过稳定化和均等化机制影响物种共存。本文首先介绍了叶片病原真菌驱动同种负密度制约(稳定化机制)和“生长-防御”权衡(均等化机制)促进物种共存的证据, 并阐述了在群落水平抑制叶片病原真菌后物种丰富度的变化。随后, 本文归纳了该领域研究中的主要挑战: 叶片病原真菌在驱动物种共存过程中相较于环境因子的可能重要性更低、部分叶片病原真菌较弱的宿主专一性无法起到维持物种共存的作用, 以及控制叶片病原真菌过程中各种方法均有一定局限性等问题。最后, 本文论述了该研究领域未来的主要方向: 不同土壤养分/气候变化条件下叶片病原真菌如何影响物种共存、叶片病原真菌与其他高营养层次生物类群的交互作用及其对物种共存的贡献、基于系统发育推断叶片病原真菌对植物群落构建的影响、将叶片病原真菌生活史类型纳入病原真菌影响植物物种共存的相关研究中。

关键词: 物种共存, 物种多样性, 叶片病原真菌, 负密度制约, 稳定化机制, 均等化机制

Abstract

Background & Aims: Understanding how species coexist is a core question in community ecology that has long intrigued ecologists. Under the contemporary coexistence theory, the mechanisms that promote species coexistence are categorized into stabilizing and equalizing mechanisms. However, we know little about how foliar fungal pathogens affect stabilizing and equalizing mechanisms, despite being abundant and important.

Progresses & Challenges: In this review, we briefly introduce the experimental evidence of how foliar fungal diseases promote plant species coexistence through conspecific negative density dependence (via stabilizing mechanisms) and growth-defense tradeoffs (via equalizing mechanisms), as well as how community-level foliar fungal pathogen removal affects plant species richness. We then discuss controversies surrounding this topic. These controversies mainly focused on the relative importance of foliar fungal pathogens in driving species coexistence compared to other factors, host specificity of foliar fungal pathogens, and methodological issues in related studies.

Prospects: We highlight some open questions for future studies on this topic, including how climate change and soil nutrients affect the relationship between foliar fungal pathogens and species coexistence, the interactions between foliar fungal pathogens and other high trophic biomes impact upon species coexistence, inferring the effects of foliar fungal pathogens on plant community assembly based on phylogeny, and incorporating plant life histories into related studies.

Key words: species coexistence, species diversity, foliar fungal pathogens, negative density dependence, stabilizing mechanisms, equalizing mechanisms

刘向, 刘木, 肖瑶 (2023) 叶片病原真菌对植物物种共存的影响: 进展与挑战. 生物多样性, 31, 22525. DOI: 10.17520/biods.2022525.

Xiang Liu, Mu Liu, Yao Xiao (2023) The effect of foliar fungal pathogens on plant species coexistence: Progress and challenges. Biodiversity Science, 31, 22525. DOI: 10.17520/biods.2022525.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2022525

https://www.biodiversity-science.net/CN/Y2023/V31/I2/22525

图/表 2

图1 叶片真菌病害影响植物物种共存的理论框架。图中直线表示植物每员增长率与同种密度之间的关系, 直线的斜率表示同种负密度制约强度, 其中蓝线表示对照, 黄线表示叶片病原真菌移除处理。对任意植物物种, 可能存在以下情况: (A)叶片真菌病害是驱动同种负密度制约的重要因子, 则叶片病原真菌移除处理会导致直线的斜率更加平缓; (B)叶片真菌病害可以降低所有同种密度条件下的宿主植物适合度, 则叶片病原真菌移除处理会导致直线在y轴的截距升高; (C)叶片真菌病害既是驱动同种负密度制约的重要因子, 也能够降低宿主植物适合度, 则叶片病原真菌移除处理会导致直线的斜率更加平缓且其在y轴的截距升高。图中, a1是对照情景(蓝线)下的截距, Δa是对照和叶片病原真菌移除处理(黄线)之间截距的差异, b1是对照情景下的回归系数, Δb是对照和叶片病原真菌移除处理之间回归系数的差异。修改自Levine和HilleRisLambers (2009)和Liu等(2022)。

Fig. 1 A framework for how foliar fungal diseases affect plant species coexistence, showing how plant per capita growth rate changes with conspecific host density and with foliar fungal pathogens removal treatment (yellow) relative to control (blue). The strength of conspecific negative density dependence (CNDD) is indicated by the slope of the line showing the relationship between the per capita growth rate and conspecific density. For any given species, we expect: (A) If foliar fungal diseases generate CNDD, foliar fungal pathogens removal should lead to a shallower slope of the line; (B) If foliar fungal diseases decrease host fitness across all densities, foliar fungal pathogens removal should lead to a greater y-intercept; (C) If foliar fungal diseases generate CNDD as well as decrease host fitness, foliar fungal pathogens removal should cause both a shallower slope of the line and a higher y-intercept. Here, a1 is intercept for control (blue line), Δa is the difference in intercept between the control and foliar fungal pathogens removal treatments; b1 is the regression coefficient for control, and Δb is the difference in regression coefficient between control and foliar fungal pathogens removal treatments. Adapted from Levine & HilleRisLambers (2009) and Liu et al (2022).

表1 自然生态系统中控制群落水平叶片真菌病害方法的比较

Table 1 Comparison of methods for controlling community-level foliar fungal diseases in natural ecosystems

方法 Methods	优势 Advantages	劣势 Disadvantages	参考文献 References
喷洒杀菌剂 Spraying fungicide	简洁、高效 Simple and efficient	引入无关化学成分、非靶标效应较强 Introducing irrelevant chemicals, and strong non-target effects	Allan et al, 2010; Seabloom et al, 2017
喷洒叶片病原真菌繁殖体 Spraying foliar fungal propagules	避免引入无关化学成分 Avoiding introducing irrelevant chemicals	并非所有叶片病原真菌均能够获得繁殖体 Not all foliar fungal pathogens are able to obtain propagules	Peters & Shaw, 1996
病叶移除 Diseased leaves removal	避免引入无关化学成分 Avoiding introducing irrelevant chemicals	费时费力、移除叶片对植物群落产生扰动 Time-consuming and labor-intensive, and the leaves removal disturbs the plant community	-

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